GIFT  OF 


UNIVERSITY  OF  CALIFORNIA  PUBLICATIONS 

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ZOOLOGY 

Vol.  16,  No.  17,  pp.  293-348,  plates  24-26  June  23,  1916 


DISTRIBUTION  OF  THE  LAND  VERTEBRATES 
OF  SOUTHEASTERN  WASHINGTON 


BY 

LEE   RAYMOND    DICE 


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8.  Distribution  of  River  Otters  in  California,  with  Description  of  a  New 

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311-316.     November,  1914 .05 

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January,  1915  .05 


DISTRIBUTION  OF  THE  LAND  VERTEBRATES 
OF  SOUTHEASTERN  WASHINGTON 


A  THESIS  SUBMITTED  IN  PARTIAL  SATISFACTION  OF 
THE  REQUIREMENTS  FOR  THE  DEGREE  OF 

DOCTOR  OF  PHILOSOPHY 
AT  THE  UNIVERSITY  OF  CALIFORNIA 


BY 
LEE  RAYMOND  DICE 


MAY,  1915 


UNIVERSITY  OF  CALIFORNIA   PUBLICATIONS 

IN 

ZOOLOGY 

Vol.  16,  No.  17,  pp.  293-348,  plates  24-26  June  23,  1916 


DISTRIBUTION  OF  THE  LAND  VERTEBRATES 
OF  SOUTHEASTERN  WASHINGTON 

BY 

LEE  EAYMOND  DICE 


CONTENTS 

PAGE 

Introduction  293 

Topography  294 

Geological  history  296 

Methods  of  study  employed 296 

Faunas  and  associations  of  the  region  298 

Columbia  Basin  sagebrush  faunal  area  and  fauna  300 

Columbia  Basin  prairie  faunal  area  and  fauna  305 

Blue  Mountain  faunal  area  and  fauna  314 

Climate 322 

Influence  of  artificial  conditions 326 

Comparison  with  other  schemes  of  ecological  distribution  327 

Zoogeographical  position  of  southeastern  Washington  329 

Life-zones  of  southeastern  Washington  331 

Comparison  of  the  different  systems  of  considering  distribution  332 

The  zoogeographical  system  332 

The  life-zone  system  333 

The  ecological  method  339 

Summary   340 

Literature  cited  ..  ..  341 


INTRODUCTION 

In  the  following  paper  the  local  and  geographical  distribution  of 
the  land  vertebrates  of  Walla  Walla  and  Columbia  counties,  south- 
eastern Washington,  is  considered.  The  region  selected  is  of  consider- 
able geographical  extent.  It  is  especially  adapted  to  the  present  study 
because  it  possesses  a  wide  range  of  climatic  conditions,  giving  a 
variety  of  habitats.  In  the  presentation  of  the  facts  the  endeavor  is 
made  to  show  how  the  consideration  of  local  distribution  may  be  com- 
bined with  the  consideration  of  geographical  distribution. 


318611 


fiK<       :K5.*:w 

294  University  of  California  Publications  in  Zoology        ITOL- 16 

The  dependence  of  animals  on  their  habitats  does  not  seem  to  have 
been  sufficiently  taken  into  account  in  the  study  of  distribution.  Many 
species  are  closely  restricted  to  certain  kinds  of  habitats.  If  a  particu- 
lar kind  of  habitat  does  not  occur  in  a  region  the  species  of  animals 
restricted  to  it  must  also  be  absent.  This  applies  to  the  habitats  of 
great  extent  as  well  as  to  the  habitats  which  always  occur  in  limited 
areas.  We  should  not  expect  to  find  typically  desert  animals  except 
in  a  desert  habitat.  In  order  to  explain  distribution  we  should  give 
much  attention  to  the  causes  producing  the  different  kinds  of  habitats. 

Usually  the  fauna  of  each  geographical  region  has  been  studied  as 
a  whole  and  whole  faunas  have  been  compared  together  without  regard 
to  differences  in  the  habitats  of  the  component  species.  It  seems  that 
such  a  method  cannot  lead  to  the  most  exact  knowledge  of  the  factors 
limiting  the  distribution  of  animals  on  the  continent  of  North  America 
or  of  the  relation  in  origin  of  particular  species  or  faunas. 

In  order  to  show  the  relation  of  the  faunal  areas  of  southeastern 
Washington  to  the  faunal  divisions  of  North  America,  the  position  of 
the  region  in  the  zoogeographical  system  of  considering  distribution 
and  in  the  life-zone  system  is  determined  as  closely  as  possible.  Finally, 
a  critical  comparison  is  made  of  the  several  methods  of  studying  animal 
distribution  which  are  at  present  in  use. 

The  latter  part  of  the  field-work  and  the  preparation  of  the  results 
have  been  supervised  by  Professor  S.  J.  Holmes,  who  has  given  much 
valuable  criticism.  I  am  also  indebted  to  Dr.  Joseph  Grinnell  for 
advice  and  criticism.  Professor  J.  C.  Merriam  furnished  most  of  the 
notes  on  the  geological  history  of  the  region. 


TOPOGRAPHY 

The  topographical  features  of  southeastern  Washington  have  been 
described  in  detail  by  Russell  (1897,  pp.  14-28).  The  western  part  of 
Walla  Walla  County  has  in  general  a  low  relief  and  is  largely  formed 
of  several  "Flats."  However,  south  of  the  Walla  Walla  River  in  this 
region  there  is  a  high  range  of  hills. 

East  of  Lamar  the  hills  become  somewhat  abruptly  higher  and  the 
prairie  region  of  eastern  Walla  Walla  County  and  western  Columbia 
County  is  characterized  by  high,  rolling  hills.  The  hills  are  steepest  on 
the  northeast  slopes  and  these  northern  hillsides  are  damper  and  colder 
and  support  a  more  luxuriant  growth  of  vegetation  than  the  southern 
slopes.  Along  the  northern  boundary  of  Walla  Walla  and  Columbia 


1916]        Dice:  Land  Vertebrates  of  Southeastern  ^Washington          295 


296  University  of  California  Publications  in  Zoology        tv°L- 16 

counties  the  Snake  River  has  cut  a  deep  canon,  reaching  in  places  to 
a  depth  of  1500  feet  or  more. 

The  Blue  Mountains  in  southeastern  Columbia  County  form  a 
region  of  steep  slopes  and  rather  sharp  ridges,  with  an  elevation  of 
4000  to  6000  feet.  The  canons  are  deep  and  the  valleys  narrow. 

GEOLOGICAL  HISTORY 

In  early  Miocene  time  the  greater  portion  of  eastern  Washington 
and  also  a  portion  of  eastern  Oregon  and  Idaho  were  flooded  by  a 
series  of  lava  flows.  These  Columbia  Lavas  have  a  thickness  in  places 
of  probably  over  4000  feet.  All  the  visible  rocks  of  Walla  Walla  and 
Columbia  counties  are  basalts  probably  formed  at  the  time  of  these 
lava  flows.  The  canon  walls  of  Snake  River  indicate  at  least  ten  suc- 
cessive flows,  between  some  of  which  sufficient  time  elapsed  for  soil  to 
form  and  forests  to  grow. 

Volcanic  outbursts  continued  during  the  Middle  Miocene  and  much 
volcanic  ash  was  thrown  out.  In  eastern  Oregon  extensive  lake  de- 
posits, the  Mascall  beds,  were  formed  at  that  time. 

The  climate  of  the  region  in  the  period  succeeding  the  lava  flows 
must  have  been  more  moist  than  at  present.  Before  the  middle  of  the 
Pliocene,  however,  the  Cascade  Mountains  had  been  elevated  and  these 
mountains  robbed  the  winds  from  the  ocean  of  their  moisture  long 
before  they  reached  eastern  Washington.  A  dry  period  necessarily 
ensued. 

The  time  of  the  uplifting  of  the  Blue  Mountains  is  not  certainly 
known,  but  their  present  elevation  had  probably  been  reached  before 
the  beginning  of  the  Pliocene. 

In  Pleistocene  time  the  presence  of  glaciers  in  northern  Washington 
seems  to  have  caused  a  return  of  moist  conditions  over  the  region.  At 
that  time  also  the  gorge  of  the  Columbia  River  through  the  Cascade 
Mountains  seems  to  have  been  blocked,  resulting  in  the  formation  of 
a  large  lake,  Lake  Lewis,  which  covered  a  large  part  of  the  Columbia 
Basin.  In  Walla  Walla  County  this  lake  probably  extended  eastward 
as  far  as  Eureka,  but  no  evidences  of  it  east  of  this  point  have  been 
found. 

METHODS  OF  STUDY  EMPLOYED 

The  principal  aim  in  the  present  study  has  been  to  determine  the 
vertebrate  associations  of  the  region.  The  wide  range  of  climatic  and 
vegetational  conditions  which  occurs  gives  a  very  good  opportunity  to 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          297 

study  the  extent  to  which  the  different  species  are  restricted  to  par- 
ticular habitats.  The  habitats  have  been  described  in  some  detail  in 
order  to  facilitate  comparison  with  the  habitats  of  other  regions. 

The  attempt  has  been  made  to  study  and  describe  the  habitats  and 
associations  in  as  nearly  their  native  and  normal  conditions  as  pos- 
sible, but  for  many  habitats  it  has  been  difficult  to  find  a  sufficient  area 
retaining  anything  like  the  original  conditions. 

While  many  of  the  associations  here  considered  could  be^easily  sub- 
divided or  several  could  perhaps  be  combined,  yet  it  is  believed  that 
the  present  arrangement  is  the  most  useful  one  for  the  study  of  verte- 
brate distribution  in  the  region.  A  region  is  divided  into  associations 
as  a  matter  of  convenience  in  studying  and  describing  the  environ- 
ments and  the  habitat  preferences  of  the  animals.  To  increase  the 
number  of  associations  unduly  would  destroy  the  convenience  of  use 
for  which  the  classification  is  made,  while  to  lump  the  associations 
might  obscure  important  facts.  It  cannot  be  hoped  that  the  relative 
abundance  of  the  species  in  the  different  associations  as  here  given  is 
fully  accurate,  for  the  observations  on  which  the  results  are  founded 
are  known  in  many  cases  to  be  too  few  in  number. 

In  order  to  show  the  relative  abundance  of  each  species  in  the  dif- 
ferent habitats  it  has  been  desirable  to  use  a  system  of  nomenclature 
modified  from  that  used  by  Grinnell  (1914,  p.  67) .  As  here  considered, 
the  relative  abundance  of  each  species  in  the  different  habitats  of  the 
same  area  is  compared.  No  attempt  is  made  to  compare  the  relative 
abundance  of  a  species  in  the  different  faunal  areas.  When  a  species 
occurs  in  only  one  habitat  in  an  area,  this  is  designated  the  exclusive 
habitat.  If  the  species  occurs  in  more  than  one  habitat  in  the  area,  the 
habitat  in  which  it  occurs  most  abundantly  is  said  to  be  its  major 
habitat  and  all  others  are  said  to  be  minor  habitats.  If  the  data  are 
insufficient  to  determine  the  major  habitat,  the  term  reported  is  used 
to  refer  to  each  habitat  in  which  the  species  is  known  to  occur.  The 
relative  abundance  in  the  different  habitats  has  been  determined  on 
the  basis  of  the  comparative  number  of  individuals  actually  observed 
or  trapped  in  each. 

Active  observation  of  the  birds  in  the  region  near  Prescott  was 
begun  by  the  author  in  December,  1904,  and  attention  was  later 
directed  to  the  other  vertebrates.  The  observations  have  been  much 
interrupted  by  the  conflicting  claims  of  other  duties  and  by  prolonged 
absences  from  the  region.  In  the  summer  of  1914  ten  days  were  spent 
in  the  region  near  Wallula,  a  shorter  excursion  was  made  to  Lyon's 


298  University  of  California  Publications  in  Zoology        tv°L- 16 

Ferry  on  the  Snake  River,  and  three  weeks  were  spent  in  the  Blue 
Mountains  in  very  intensive  study.  Notes  published  by  others  bearing 
on  the  distribution  of  the  vertebrates  of  the  region  have  been  used 
whenever  possible.  Mr.  S.  H.  Lyman  of  Dayton  has  kindly  allowed 
the  use  of  his  unpublished  notes  on  the  birds  of  the  region.  It  is  hoped 
that  the  full  accounts  of  the  distribution  of  each  species  of  vertebrates 
in  the  region,  which  were  prepared  in  working  up  the  present  paper, 
may  soon  be  published. 

Great  care  has  been  taken  to  obtain  accuracy  in  the  specific  identifi- 
cations. Specimens  have  been  secured  when  possible,  except  of  easily 
recognized  forms.  In  a  few  cases  the  subspecific  identification  is  based 
on  the  geographical  distribution  as  given  by  recognized  authorities. 
Dr.  Joseph  Grinnell  has  checked  a  large  number  of  identifications  of 
mammals  and  birds.  The  identification  of  the  reptiles  and  amphibians 
is  due  to  Mr.  Charles  L.  Camp. 

The  botanical  names  used  have  been  taken  from  Piper  (1906)  ex- 
cept in  a  few  cases  where  other  names  seemed  more  desirable.  Speci- 
mens of  many  of  the  more  important  species  of  plants  were  collected. 
Dr.  H.  M.  Hall  identified  a  number  of  specimens  and  Dr.  H.  S.  Yates 
determined  several  grasses. 


FAUNAS  AND  ASSOCIATIONS  OF  THE  REGION 

In  southeastern  Washington  three  prevailing  types  of  vegetation 
may  be  recognized.  Along  the  Columbia  River  there  is  an  area  where 
sagebrush  is  the  dominant  plant ;  further  east  is  a  region  where  bunch- 
grass  forms  the  most  prominent  part  of  the  natural  vegetation;  and 
the  Blue  Mountains  are  largely  covered  by  conifer  forests.  Correlated 
with  these  differences  in  vegetation  there  are  important  differences  in 
the  species  of  vertebrates  found  in  each  of  these  districts.  The  as- 
semblage of  species  found  in  each  such  region  delimited  by  climatic 
features  may  be  called  a  fauna,  and  the  region  itself  may  be  called  a 
faunal  area,  or,  more  simply,  an  area.  The  term  *  *  area ' '  as  here  used 
refers  to  the  whole  of  any  geographical  district  where  a  particular 
type  of  vegetation  is  dominant  and  includes  all  of  the  habitats  in  such 
a  district. 

Each  faunal  area  is  made  up  of  several  different  kinds  of  habitats 
and  each  habitat  shelters  a  different  association  of  vertebrates.  Some 
habitats  and  associations  in  the  different  faunal  areas  are  very  similar, 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          299 

but  in  the  present  paper  an  association  is  not  considered  to  extend  be- 
yond the  limits  of  a  single  faunal  area. 

The  lines  separating  the  different  faunal  areas  are  not  sharp.  Prob- 
ably the  best  criterion  for  characterizing  faunal  areas  is  the  dominance 
of  particular  habitats.  It  is  evident  that  in  passing  from  one  area  to 
another  a  situation  will  be  met  where  the  dominant  habitat  of  one  area 
will  equal  in  extent  the  dominant  habitat  of  the  other  area.  It  is  at 
this  point  that  the  line  separating  the  two  must  be  drawn.  -Each  area 
will  usually  show  at  its  edges  some  development  of  the  dominant 
habitats  of  the  adjacent  areas.  In  many  cases  a  dominant  habitat 
from  one  area  may  recur  as  a  subdominant  habitat  throughout  an 


Fig.  B.  Map  of  Walla  Walla  and  Columbia  counties,  southeastern  Wash- 
ington, showing  the  extent  of  each  of  the  three  faunal  areas,  Columbia  Basin 
sagebrush  area,  Columbia  Basin  prairie  area,  and  Blue  Mountain  area. 


adjacent  area.     However,  it  is  best  to  consider  each  area  in  sections 
where  it  is  typically  developed  and  not  along  its  edges. 

Lists  of  the  characteristic  species  of  each  of  the  faunas  are  given. 
Each  list  includes  those  native  breeding  species  which  in  southeastern 
Washington  are  definitely  known  from  only  one  faunal  area.  Lists 
are  also  given  of  the  species  making  up  the  different  associations  of 
each  fauna.  As  here  given  the  lists  record  all  the  forms  noted  in  each 
of  the  corresponding  habitats.  Trapping  records  are  included  in  some 
instances  to  show  the  relative  abundance  of  some  of  the  smaller  mam- 
mals. Unless  otherwise  noted,  only  the  results  of  the  first  night's 
trapping  on  any  trap  line  are  included.  Traps  are  usually  set  from 
five  to  ten  yards  apart  and  in  a  continuous  line. 


300  University  of  California  Publications  in  Zoology        [VOL.  16 

COLUMBIA  BASIN  SAGEBRUSH  FAUNAL  AREA  AND  FAUNA 
Habitats  and  Associations: 

Sagebrush.  Water-margin. 

Rocky-slope.  Aquatic. 

Willow.  Aerial. 

The  Columbia  Basin  sagebrush  area  is  characterized  by  the 
dominance  of  the  sagebrush  habitat.  This  is  found  well  developed  in 
the  western  end  of  Walla  Walla  County  near  the  Columbia  and  Snake 
rivers.  Sagebrush  extends  up  the  valley  of  the  Walla  Walla  River  to 
the  neighborhood  of  the  town  of  Touchet  or  a  little  above  this.  It 
also  extends  up  Snake  River  for  some  distance,  but  the  exact  limits 
are  not  known.  Sagebrush  is  not  dominant  south  of  the  Snake  River 
at  Lyon  '&  Ferry,  so  the  sagebrush  area  does  not  extend  eastward  that 
far.  Between  the  Walla  Walla  and  Snake  rivers  the  sagebrush  area 
extends  eastward  a  number  of  miles,  but  as  the  land  rises  the  sagebrush 
gradually  gives  place  to  the  prairie.  The  change  is  very  gradual  and 
no  abrupt  line  of  demarcation  can  be  drawn.  The  typical  sagebrush 
area  probably  does  not  extend  eastward  from  the  Columbia  River  more 
than  about  ten  miles. 

There  are  some  rocky  slopes  in  all  the  higher  hills  of  the  sagebrush 
area  and  along  the  streams  there  are  numerous  basaltic  bluffs.  Along 
the  Walla  Walla  River  there  is  a  narrow  growth  of  willows,  but  along 
the  Columbia  and  Snake  rivers  in  western  Walla  Walla  County  there 
are  almost  no  trees,  and  brush  is  developed  in  only  a  few  places,  so 
that  along  these  streams  the  willow  habitat  appears  only  in  isolated 
patches. 

CHARACTERISTIC  SPECIES  OF  THE  COLUMBIA  BASIN  SAGEBRUSH  FAUNA 

Scaphiopus  hammondii  hammondii.  Lanius  ludovicianus  excubitorides. 

Sceloporus  graciosus.  Onychomys  leucogaster  fuscogriseus. 

Phrynosoma  douglassii  douglassii.  Perognathus  parvus  parvus. 

Centrocercus  urophasianus.  Perognathus  lordi  columbianus. 

Amphispiza  nevadensis  nevadensis.  Perodipus  ordii  columbianus. 

The  Columbia  Basin  sagebrush  fauna  is  characterized  by  the 
presence  of  a  considerable  number  of  species  nearly  all  of  which  are 
specially  adapted  to  semi-desert  conditions  and  are  inhabitants  of  the 
sagebrush  habitat. 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington 


301 


SAGEBEUSH  HABITAT  AND  ASSOCIATION  (SAGEBRUSH  AEEA) 


Exclusive : 

Sceloporus  graciosus. 
Phrynosoma  douglassii  douglassii. 
Speotyto  cunicularia  hypogaea — 

summer. 

Sturnella  neglecta — summer. 
Chondestes  grammacus  strigatus — 

summer. 
Amphispiza  nevadensis  nevadensis 

— summer. 

Major : 

Scaphiopus  hammondii  hammondii. 
Chordeiles  virginianus  hesperis — 
summer. 


Lanius  ludovicianus  excubitorides 

— summer. 
Canis  latrans  lestes. 
Taxidea  taxus  neglecta. 
Perognathus  parvus  parvus. 
Perodipus  ordii  columbianus. 
Citellus  townsendii. 
Lepus  californicus  wallawalla. 


Sylvilagus  nuttallii  nuttallii. 


Minor: 

Crotalus  oregonus. 

Oxyechus  vociferus  vociferus — 

summer. 
Zenaidura  macroura  marginella — 

summer. 


Pica  pica  hudsonia — resident. 
Icterus  bullocki — summer. 
Euphagus  cyanocephalus — summer. 
Thomomys  columbianus. 


Reported : 

Pedioecetes    phasianellus    colum- 
bianus— resident. 

Centrocercus  urophasianus — 
resident. 

Falco  sparverius  sparverius — 
summer. 


Asio  flammeus — summer. 
Astragalinus  tristis  pallidus — 

summer. 
Onychomys  leucogaster  fuscogriseus. 


The  sagebrush  habitat,  where  it  was  studied  three  miles  east  of 
Wallula,  is  not  entirely  homogeneous.  The  dominant  plant  is  the 
common  sagebrush  (Artemisia  tridentata).  Commonly  mixed  with 
this  are  two  species  of  rabbit  brush  (Chrysotkamnus  viscidiflorus  and 
Chrysothamnus  nauseosus  graveolens).  In  places  one  or  other  of 
these  shrubs  may  be  more  abundant  than  the  sagebrush.  The  hop  sage 
(Grayia  spinosa)  and  the  antelope  brush  (Kunzia  tridentata}  occur  in 
lesser  abundance.  In  sandy  areas  a  cactus  (Opuntia  polyacantha)  is 
often  found.  The  wheat  bunchgrass  (Agropyron  spicatum)  is  found 
very  sparingly,  but  under  native  conditions  was  evidently  much  more 
abundant  than  at  present.  Where  there  has  been  extensive  pasturage 
and  tramping  by  stock  the  yarrow  (Achillea  mille folium  lanulosa)  is 
common. 


302  University  of  California  Publications  in  Zoology        tv°L- 16 

The  soil  in  the  sagebrush  habitat  is  light  and  sandy  and  being  sub- 
jected to  high  winds  often  drifts,  and  areas  of  drifting  sand  are  com- 
mon. The  sand  heaps  up  about  the  various  shrubs,  forming  small 
dunes.  Being  continually  shifting  it  would  not  seem  to  be  a  good  place 
for  ground-dwelling  animals  to  make  their  homes.  There  are  small 
areas  where  there  are  no  shrubs  or  plants  but  only  drifting  sand, 
which  in  some  places  near  the  larger  rivers  forms  good-sized  dunes. 
Over  large  areas  covered  by  sagebrush  the  sand  is  packed  and  is  being 
eroded  by  the  wind.  In  these  places  the  sand  is  removed  as  soon  as  it 
is  loosened  so  that  little  loose  sand  is  present.  By  the  erosion  small 
sand  bluffs  are  sometimes  exposed. 

The  sagebrush  association  is  represented  by  a  considerable  number 
of  species,  most  of  which  are  characteristic  of  semi-desert  conditions. 
A  few  birds,  which  breed  along  the  streams,  forage  out  a  considerable 
distance  into  the  sagebrush. 

Trapping  in  sagebrush  three  miles  east  of  Wallula  on  the  nights  of 
June  10,  June  12,  and  June  17,  1914,  produced  a  total  of  1  Onychomys 
leucogaster  fuscogriseus,  6  Perognathus  parvus  parvus.  and  6  Pero- 
dipus  ordii  columbianus.  On  these  nights  there  were  61,  66,  and  61 
traps  used  respectively.  This  gives  a  total  of  188  "trap-nights" 
(Grinnell,  1914,  p.  92).  Most  of  the  traps  were  "  out-o-sight "  mouse 
traps,  but  5  or  6  were  rat  traps. 

EOCKY-SLOPE  HABITAT  AND  ASSOCIATION  (SAGEBRUSH  AEEA) 

Exclusive : 

Salpinetes  obsoletus  obsoletus — summer. 

Major : 

Crotalus  oregomis. 

Minor: 

Peromyscus  maniculatus  gambelii.  Sylvilagus  nuttallii  nuttallii. 

Reported  : 

Buteo  borealis  calurus — summer  Neotoma  cinerea  occidentalis. 

The  rocky-slope  habitat  is  made  up  of  the  slopes  covered  by  broken 
rock  and  of  the  exposures  of  solid  basalt  and  their  talus  slopes.  Some 
vegetation  is  usually  found  in  the  soil  among  the  rocks  and,  because 
the  basalt  rapidly  decomposes,  there  is  a  tendency  for  plants  to  in- 
crease rapidly  in  numbers.  The  vegetation  usually  agrees  in  character 
with  that  of  the  surrounding  country.  Sagebrush  (Artemisia  tri- 
dentata)  and  wheat  bunchgrass  (Agropyron  spicatum)  usually  both 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington 


303 


occur  and  one  or  the  other  is  dominant,  depending  on  whether  the 
region  is  dominated  by  sagebrush  or  bunchgrass.  In  general  the 
habitat  is  strikingly  arid. 

The  species  of  the  rocky-slope  association  are  few  in  number  and 
represent  species  which  in  general  show  a  fondness  for  the  neighbor- 
hood of  rocks. 

In  ten  traps  set  among  rocks  on  a  steep  hillside  three  miles  south- 
east of  Wallula  one  Peromzyscus  maniculatus  gambelii  was_taken  on 
June  16,  1914. 


WILLOW  HABITAT  AND  ASSOCIATION  (SAGEBRUSH  AREA) 


Exclusive : 

Colinus  virginianus  virginianus — 

resident. 

Asio  wilsonianus — resident. 
Otus  asio  macfarlanei — summer. 
Colaptes  cafer  collaris — summer. 
Corvus  brachyrhynchos  hesperis — 

summer. 
Molothrus  ater  artemisiae — 

summer. 
Melospiza  melodia  merrilli — 

resident. 

Zamelodia  melanocephala — summer. 
Passerina  amoena — summer. 

Major  : 

Zenaidura  macroura  marginella: — 

summer. 

Tyrannus  tyrannus — summer. 
Tyrannus  verticalis — summer. 
Pica  pica  hudsonia — resident. 
Icterus  bullocki — summer. 

Minor: 

Thamnophis  elegans. 

Reported : 

Pituophis  catenifer  catenifer. 


Dendroica  aestiva  aestiva — 

summer. 

Icteria  virens  longicauda — summer. 
Dumetella  carolinensis — summer. 
Penthestes  atricapillus  septentrion- 

alis — resident. 
Planesticus     migratorius     propin- 

quus — summer. 
Sorex  vagrans  dobsoni. 
Reithrodontomys  megalotis 

nigrescens. 
Mus  musculus  musculus. 


Euphagus  cyanocephalus — 

summer. 

Mephitis  occidentalis  major. 
Peromyscus  maniculatus  gambelii. 
Thomomys  columbianus. 


Sylvilagus  nuttallii  nuttallii. 
Erethizon  epixanthum  epixanthum. 


The  timber  found  along  the  Walla  Walla  River  near  Wallula  is 
made  up  largely  of  willows  (Salix)  of  several  species.  Cottonwoods 
(Populus  trichocarpa)  are  rare  and  so  are  many  of  the  shrubs  which 
grow  along  the  streams  nearer  the  Blue  Mountains.  The  habitat  is 
limited  in  extent  and  does  not  usually  extend  more  than  a  few  rods 
from  the  banks  of  the  river,  when  it  ceases  abruptly  and  gives  place  to 
the  sagebrush.  The  willows  are  mostly  small  in  size  and  grow  very 


304  University  of  California  Publications  in  Zoology        [VOL.  16 

thickly  together,  forming  a  dense  thicket.  On  some  of  the  very  low 
land  near  the  river  there  are  small  meadows  where  the  willows  have 
not  been  able  to  establish  themselves  or  from  which  they  have  been 
cleared  by  man. 

The  majority  of  the  species  of  the  willow  association  are  not  found 
in  the  adjacent  sagebrush  association  and  these  two  associations  are 
very  distinct.  The  species  of  the  willow  association  in  the  sagebrush 
area  are  all  found  also  in  the  cottonwood-willow  association  of  the 
prairie  area. 

Twenty-nine  traps  set  at  the  edge  of  the  willows  along  the  Walla 
Walla  River  three  miles  east  of  Wallula  took,  on  June  13,  1914,  7 
Reithrodontomys  megalotis  nigrescens,  8  Peromyscus  maniculatus  gam- 
belii,  and  6  Mus  musculus  musculus.  On  June  15,  the  third  day's  trap- 
ping in  this  trap-line,  one  Sorex  vagrans  dobsoni  was  taken,  and  also 
several  mice.  It  is  evident  that  small  mammals  are  much  more  numer- 
ous in  these  willows  than  in  other  habitats  of  the  area.  This  is  true  of 
birds  also,  for  more  were  seen  here  than  elsewhere. 

WATER-MARGIN  HABITAT  AND  ASSOCIATION  (SAGEBRUSH  AREA) 

Major  : 

Thamnophis  elegans.  Oxyechus  vociferus  vociferus — 

summer. 

Minor: 

Scaphiopus  hammondii  hammondii. 

Reported  : 

Rana  pipiens  brachycephala.  Agelaius  phoeniceus  neutralis — 

summer. 

The  water-margin  habitat  comprises  the  shores  of  streams,  irrigat- 
ing ditches,  and  lakes.  Only  a  few  seepage  lakes  occur  in  western 
Walla  Walla  County  and  streams  and  irrigating  ditches  are  not  numer- 
ous. There  is  a  considerable  extent  of  barren  sandy  and  gravelly  shore 
along  the  Columbia  and  Snake  rivers.  Little  study  was  made  of  the 
life  of  those  places.  Along  the  Walla  Walla  River  the  water-margin 
habitat  is  a  very  narrow  strip  between  the  river  and  the  growth  of 
willows.  A  few  bars  of  mud  and  gravel  occur,  but  these  are  not  ex- 
tensive. 

In  the  sagebrush  area  the  water-margin  association  is  not  very  im- 
portant and  is  made  up  of  only  a  few  characteristic  species. 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          305 

AQUATIC  HABITAT  AND  ASSOCIATION  (SAGEBRUSH  AREA) 

Beported  : 

Ondatra  zibethicus  osoyoosensis. 

The  aquatic  association  as  here  considered  includes  the  animals 
which  inhabit  the  open  water  of  the  streams  away  from  the  proximity 
of  the  shores.  Vertebrate  .members  of  the  association  other  than  the 
fishes  are  few  in  the  region.  The  Columbia  and  Snake  rivers_are^both 
swift  and  during  most  of  the  year  carry  much  mud  and  sand.  The 
lower  part  of  the  Walla  Walla  River  is  much  more  sluggish  and  there 
are  a  number  of  quiet  pools.  However,  there  is  little  aquatic  vegeta- 
tion and  few  aquatic  insects,  and  the  habitat  does  not  appear  very 
suitable  for  the  higher  vertebrates. 

AERIAL  HABITAT  AND  ASSOCIATION  (SAGEBRUSH  AREA) 

Minor: 

Chordeiles  virginianus  hesperis —  Tyrannus  tyrannus — summer, 

summer.  Tyrannus  verticalis — summer. 

Reported: 

Tachycineta  thalassina  lepida — summer. 

The  aerial  association  is  considered  to  be  made  up  of  those  animals 
which  feed  in  the  air.  Most  birds  fly  about  in  the  air  more  or  less, 
but  the  association  should  be  limited  to  those  species  which  carry  on  a 
vital  activity  in  the  habitat. 


COLUMBIA  BASIN  PRAIRIE  FAUNAL  AREA  AND  FAUNA 

Habitats  and  Associations  : 

Bunchgrass.  Water-margin. 

Rocky-slope.  Aquatic. 

Cottonwood-willow.  Aerial. 

The  bunchgrass  habitat  is  dominant  over  most  of  Walla  Walla 
County  and  the  western  part  of  Columbia  County.  In  typical  parts  of 
the  area  it  covers  the  entire  region  with  the  exception  of  small  areas 
of  rocky  slopes  and  the  small  amount  of  surface  occupied  by  the 
streams  and  their  adjacent  habitats.  Sagebrush  is  well  developed  in 
local  areas  on  the  flats  to  the  west  of  Lamar  and  in  a  few  places  in  the 
canons  near  Snake  River,  but  in  the  typical  part  of  the  prairie  area 
sagebrush  is  clearly  subordinate  to  the  bunchgrass  and  does  not  form 


306 


University  of  California  Publications  in  Zoology        [VoL- 16 


a  distinct  habitat.  Near  Nine-mile,  on  the  Walla  Walla  River,  the 
valley  is  definitely  semi-desert,  and  sagebrush  extends  up  the  side 
canons,  but  the  vegetation  of  the  higher  land  is  dominated  by  bunch- 
grass.  Bunchgrass  is  also  dominant  on  the  upper  parts  of  the  range 
of  hills  south  of  the  Walla  Walla  River.  Yellow  pines  invade  the 
prairie  from  the  Blue  Mountains,  coming  down  along  the  north  and 
northeastern  hillsides  and  appearing  in  the  bottoms  of  the  canons  in 
the  foothills. 

Along  Snake  River  the  region  is  drier  than  at  Prescott  and  the 
rocks  lie  closer  to  the  surface.  Along  the  canons  in  that  region  there 
are  high  basaltic  bluffs  and  many  rocky  slopes.  Toward  the  Blue 
Mountains  also  there  are  numerous  outcroppings  of  rocks,  but  over 
most  of  the  prairie  area  rocks  are  rarely  found  at  the  surface  of  the 
ground. 

Streams  are  not  numerous  in  the  prairie  area.  There  is  a  growth 
of  willows  and  other  deciduous  trees  and  shrubs  along  the  smaller 
streams,  but  along  Snake  River  there  are  few  native  trees  or  shrubs, 
so  that  no  cottonwood-willow  habitat  is  formed  along  this  stream. 


CHARACTERISTIC  SPECIES  OF  THE  PRAIRIE  FAUNA 


Aneides  iecanus. 
Hyla  regilla. 
Chrysemys  bellii. 
Actitis  macularius. 
Numenius  americanus. 
Accipiter  velox. 
Buteo  swainsoni. 
Bubo  virginianus  occidentalis. 
Dryobates  pubescens  homorus. 
Asyndesmus  lewisi. 
Archilochus  alexandri. 
Selasphorus  rufus. 
Myiochanes  richardsoni  richard- 
soni. 


Empidonax  trailli  trailli. 
Ammodramus  savannarum  bimacu- 

latus. 

Biparia  riparia. 
Stelgidopteryx  serripennis. 
Vireosylva  gilva  swainsoni. 
Geothlypis  trichas  occidentalis. 
Setophaga  ruticilla. 
Troglodytes  aedon  parkmani. 
Scapanus  orarius  schefferi. 
Mustela  arizonensis. 
Microtus  nanus  canescens. 
Perognathus  lordi  lordi. 


The  majority  of  species  known  from  the  Columbia  Basin  prairie 
area  are  known  also  from  the  sagebrush  area  or  from  the  Blue  Moun- 
tains. Of  the  species  here  given  as  unique  most  will  probably  later  be 
found  to  occur  in  the  adjacent  faunas. 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          307 


BUNCHGBASS  HABITAT  AID  ASSOCIATION  (PRAIEIE  AREA) 


Exclusive : 

Numenius  americanus — summer. 
Archibuteo  ferrugineus — summer. 
Asio  flammeus — summer. 
Speotyto  cunicularia  hypogaea — 

summer. 
Otocoris  alpestris  arcticola — 

winter. 
Otocoris  alpestris  merrilli — 

resident. 

Major  : 

Pituophis  catenifer  catenifer. 
Pedioecetes  phasianellus  colum- 

bianus — resident. 
Buteo  borealis  calurus — resident. 
Buteo  swainsoni — summer. 
Falco  mexicanus — resident. 
Sturnella  neglecta — resident. 

Minor: 

Crotalus  oregonus. 

Anas  platyrhynchos — winter. 

Ardea  herodias  treganzai — 

resident. 
Oxyechus  vociferus  vociferus — 

resident. 

Phasianus  torquatus — resident. 
Zenaidura  macroura  marginella — 

resident. 

Accipit^r  velox — resident. 
Falco  sparverius  sparverius — 

resident. 

Asyndesmus  lewisi — summer. 
Colaptes  cafer  collaris — resident. 
Tyrannus  tyrannus — summer. 
Tyrannus  verticalis — summer. 
Sayornis  sayus — summer. 
Pica  pica  hudsonia — resident. 
Molothrus  ater  artemisiae — 

summer. 

Eeported : 

Pisobia  bairdi — migrant. 


Ammodramus  savannarum  bimacu- 

latus — summer. 
Chondestes  grammacus  strigatus 

— summer. 

Spizella  breweri — summer. 
Anthus  rubescens — migrant. 
Perognathus  lordi  lordi. 
Lepus  campestris  townsendiiT~ 
Lepus  californicus  wallawalla. 

Passerculus  sandwichensis  alau- 

dinus — migrant. 
Siala  currucoides — summer. 
Canis  latrans  lestes. 
Taxidea  taxus  neglecta. 
Citellus  townsendii. 


Icterus  bullocki — summer. 
Euphagus  cyanocephalus — resident. 
Astragalinus  tristis  pallidus — 

resident. 
Spizella  passerina  arizonae — 

summer. 
Salpinctes  obsoletus  obsoletus — 

summer. 
Planesticus  migratorius  propin- 

quus — resident. 
Sialia  mexicana  occidentalis — 

migrant. 

Mustela  arizonensis. 
Mephitis  occidentalis  major. 
Reithrodontomys  megalotis 

nigrescens. 

Peromyscus  maniculatus  gambelii. 
Thomomys  columbianus. 
Citellus  columbianus  columbianus. 


Cathartes  aura  septentrionalis- 
migrant. 


The  bunchgrass  habitat  is  characterized  by  the  wheat  bunchgrass 
(Agropyron  spicatum).  With  this  are  associated  the  balsam  root 
(Balsamorhiza  sagittata),  clarkia  (Clarkia  pulchella),  Indian  bullet 
(Lithospermum  ruderale],  phlox  (Phlox  sp.)  and  several  lupines 


308  I' niversity  of  California  Publications  in  Zoology        [VOL.  16 

(Lupinus).  Common  sagebrush  (Artemisia  tridentata)  and  rabbit 
brush  (Ckrysotkamniu  viscidiflorus  and  Chrysothamnus  nauseosus 
graveolens)  occur  sometimes  in  the  bottoms  of  the  drier  ravines  and 
on  exposed  hillsides.  In  damp  situations,  in  the  bottoms  of  ravines 
or  on  north  hillsides,  the  rye  grass  (Elymus  condensatus)  forms  large 
clumps.  Where  the  land  has  been  much  pastured  the  yarrow  (Achil- 
lea  mille folium  lanulosa)  grows  abundantly.  There  are  also  a  number 
of  kinds  of  mustards  and  many  other  less  important  plants.  In  damp 
places  on  the  north  hillsides  a  few  woody  shrubs  may  be  found  and 
these  are  more  numerous  the  more  closely  the  mountains  are  ap- 
proached. The  more  important  of  these  are  the  rose  (Rosa)  and 
service-berry  (Amelanchier  ftorida}. 

Near  Prescott  almost  all  of  the  bunchgrass  hills  have  been  plowed 
and  are  planted  to  wheat  and  barley.  On  alternate  years  the  land  is 
allowed  to  lie  fallow.  The  bunchgrass  which  remains  unplowed  has 
been  heavily  pastured,  so  that  the  grass  has  been  partially  killed  out 
and  yarrow,  lupine,  and  other  weeds  have  greatly  increased. 

The  bunchgrass  association  includes  a  considerable  number  of 
plains-loving  species. 

Traps  set  in  bunchgrass  habitat  in  the  hills  two  miles  southeast 
of  Prescott  produced,  for  a  total  of  148  "trap-nights,"  on  June  27, 
July  1,  and  July  7,  1914,  five  Perognathus  lordi  lordi.  One  of  these 
trap  lines,  of  33  traps,  produced  on  the  second  day 's  trapping,  July  8, 
one  Peromyscus  maniculatus  gambelii. 

ROCKY-SLOPE  HABITAT  AND  ASSOCIATION   (PRAIRIE  AREA) 

Major : 

Crotalus  oregonus.  Salpinctes  obsoletus  obsoletus — 

summer. 

Minor: 

Pituophis  catenifer  catenifer.  Tyrannus  verticalis — summer. 

Zenaidura  macroura  marginella —  Petrochelidon  lunifrons  lunifrons 

resident.  — summer. 

Buteo  borealis  calurus — resident.  Peromyscus  maniculatus  gambelii. 

Falco  sparverius  sparverius —  Sylvilagus  nuttallii  nuttallii. 

resident. 


There  is  usually  very  little  sagebrush  growing  among  the  rocks  in 
the  rocky-slope  habitat  of  the  prairie  area.  The  wheat  bunchgrass 
(Agropyron  spicatum)  grows  abundantly  among  the  rocks  wherever 
soil  is  present.  Along  Snake  River  a  few  shrubs  grow  among  the 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington 


309 


rocks  on  the  north  slopes.  The  most  important  of  these  are  a  service- 
berry  (Amelanchier  sp.)  and  a  rose  (Rosa  sp.). 

The  characteristic  species  of  the  rocky-slope  association  are  few  in 
number.  Several  species  of  birds  find  suitable  nesting  sites  about  rock 
cliffs. 

About  the  rocks  and  rock  cliffs  near  Lyon's  Ferry  7  Peromyscus 
maniculatus  gambelii  were  taken  on  June  23  and  24,  1914,  from  115 
trap-nights. 


COTTONWOOD-WILLOW  HABITAT  AND  ASSOCIATION  (PEAIEIE  AREA) 

Exclusive : 

Aneides  iecanus. 

Bufo  columbiensis. 

Hyla  regilla. 

Perdix  perdix — resident. 

Colinus  virginianus  virginianus — 

resident. 

Bonasa  umbellns  togata — resident. 
Aceipiter  cooperi — summer. 
Astur  atricapillus  striatulus — 

winter. 

Asio  wilsonianus — resident. 
Otus  asio  macfarlanei — resident. 
Bufo  virginianus  lagophonus — 

winter. 
Bufo  virginianus  occidentalis — 

resident. 

Dryobates  villosus  orius — winter. 
Dryobates  pubescens  homorus — 

resident. 
Phloeotomus  pileatus  picinus — 

migrant. 

Archilochus  alexandri — summer. 
Selasphorus  rufus — summer. 
Cyanocitta  stelleri  annecteus — 

winter. 
Corvus  brachyrhynchos  hesperis — 

resident. 
Hesperiphona  vespertina  montana 

— winter. 

Spinus  pinus  pinus — winter. 
Passer  domesticus — resident. 
Zonotrichia  leueophrys  gambeli — 

winter. 
Spizella  monticola  ochracea — 

winter. 
Junco  hyemalis  shufeldti — winter. 


Melospiza  melodia  merrilli — 

resident. 
Passerella  iliaca  schistacea — 

summer. 

Pipilo  maculatus  curtatus — winter. 
Zamelodia  melanocephala — summer. 
Passerina  amoena — summer. 
Piranga  ludoviciana — summer. 
Bombycilla  garrula — winter. 
Vireosylva  olivacea — summer. 
Vireosylva  gilva  swainsoni — 

summer. 

Dendroica  aestiva  aestiva — summer. 
Dendroica  auduboni  auduboni — 

migrant. 

Dendroica  townsendi — migrant. 
Oporornis  tolmiei — summer. 
Geothlypis  trichas  occidentalis — 

summer. 

Icteria  virens  longicauda — summer. 
Wilsonia  pusilla  pileolata — migrant. 
Dumetella  carolinensis — summer. 
Troglodytes  aedon  parkmani — 

summer. .  , !; 

Nannus  hiemalis  pacificus — 

migrant. 

Certhia  familiaris  montana — winter. 
Sitta  carolinensis  aculeta — summer. 
Sitta  canadensis — winter. 
Penthestes  atricapillus  septentrion- 

alis — resident. 
Penthestes  gambeli  gambeli — 

winter. 
Penthestes  rufescens  rufescens — 

winter. 
Eegulus  satrapa  olivaceus— winter. 


310 


University  of  California  Publications  in  Zoology        [VOL.  16 


Kegulus  calendula  calendula — 

winter. 

Myadestes  townsendi — winter. 
Hylocichla  guttata  sequoiensis — 

summer. 


Ixoreus  naevius  naevius — migrant. 
Sorex  vagrans  dobsoni. 
Microtus  nanus  canescens. 
Mus  musculus  musculus. 
Eutamias  amoenus  amoenus. 


Major : 

Phasianus  torquatus — resident. 
Zenaidura  macroura  marginella — 

resident. 

Accipiter  velox — resident. 
Falco  sparverius  sparverius — 

resident. 
Pandion  haliaetus  carolinensis — 

summer. 

Asyndesmus  lewisi — summer. 
Colaptes  cafer  collaris — resident. 
Tyrannus  tyrannus — summer. 
Tyrannus  verticalis — summer. 
Sayornis  sayus — summer. 
Nuttallornis  borealis — migrant. 
Myiochanes  richardsoni  richard- 

soni — summer. 
Empidonax  difficilis  difficilis — 

summer. 

Empidonax  trailli  trailli — summer. 
Empidonax  wrighti — summer. 
Pica  pica  hudsonia — resident. 


Molothrus  ater  artemisiae — 

summer. 

Icterus  bullocki — summer. 
Euphagus  cyanocephalus — resident. 
Astragalinus  tristis  pallidus — 

resident. 
Spizella  passerina  arizonae — 

summer. 

Setophaga  ruticilla — summer. 
Planesticus  migratorius  propin- 

quus — resident. 
Sialia  mexicana  occidentalis — 

migrant. 

Mustela  arizonensis. 
Mephitis  occidentalis  major. 
Keithrodontomys  megalotis 

nigrescens. 

Peromyscus  maniculatus  gambelii. 
Thomomys  columbianus. 
Citellus  columbianus  columbianus. 
Sylvilagus  nuttallii  nuttallii. 


Minor: 

Eana  pipiens  brachycephala. 

Pituophis  catenifer  catenifer. 

Thamnophis  elegans. 

Ardea  herodias  treganzai — resident. 

Pedioecetes  phasianellus  colum- 
bianus— resident. 

Buteo  borealis  calurus — resident. 

Buteo  swainsoni — summer. 

Falco  mexicanus — resident. 

Streptoceryle  alcyon  caurina — 
resident. 

Agelaius  phoeniceus  neutralis — 
summer. 

Sturnella  neglecta — resident. 

Reported : 

Bascanion  constrictor  vetustum. 
Cryptoglaux  acadica  acadica — 

winter. 
Lanius  borealis — winter. 


Passerculus  sandwichensis  alau- 

dinus — migrant. 

Stelgidopteryx  serripennis — summer. 
Sialia  currucoides — summer. 
Myotis  yumanensis   (?). 
Myotis  californicus  californicus. 
Lasiurus  cinereus. 
Canis  latrans  lestes. 
Procyon  psora  pacifica. 
Mustela  vison  energumenos. 
Taxidea  taxus  neglecta. 
Citellus  townsendii. 
Castor  canadensis  canadensis. 


Seiurus  noveboracensis  notabilis — 

migrant. 
Lynx  sp. 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          311 

The  growth  of  timber  along  the  smaller  streams  of  the  prairie  area 
does  not  extend  far  from  the  banks  of  the  streams.  In  most  places 
along  the  Touchet  River  trees  do  not  naturally  grow  more  than  a 
quarter  of  a  mile  from  the  stream,  and  often  the  width  of  the  habitat 
is  much  less  than  this.  As  the  valley  of  the  Touchet  near  Fresco tt  is 
nearly  a  mile  broad  on  the  average,  it  is  evident  that  the  growth  of 
trees  and  brush  covers  only  a  portion  of  the  nearly  level  floor  of  the 
valley. 

The  most  conspicuous  plants  of  the  habitat  are  the  cottonwood 
(Populus  trichocarpa)  and  willows  (Salix)  of  several  species.  Other 
trees  and  shrubs  which  are  common  along  the  banks  of  the  Touchet 
River  near  Prescott  are  the  birch  (Betula  microphylla) ,  alder  (Alnus 
rhombifolia),  chokecherry  (Prunus  demissa},  thorn  (Crataegus  brevi- 
spina) ,  service-berry  (Amelanchier  florida),  red  osier  (Cornus  stolo- 
nifera),  and  syringa  (Philadelphus  lewisii).  Less  important  species 
are  the  cascara  sagrada  (Rhamnus  purshiana),  ninebark  (Opulaster 
pauciflorus) ,  elder  (Sambucus  glauca),  wild  cherry  (Prunus  emar- 
ginata},  snowberry  (Symphoricarpus  sp.),  and  clematis  (Clematis 
ligusticifolia) .  Roses  (Rosa  sp.)  occur  commonly,  especially  along  the 
outer  margins  of  the  timber.  The  cottonwood  often  makes  very  large 
trees  with  a  height  of  80  to  100  feet  and  with  trunks  three  to  four 
feet  in  diameter,  but  the  other  trees  are  much  smaller.  Under  the 
trees  there  is  nearly  always  a  heavy  growth  of  shrubby  underbrush. 
A  growth  of  shrubs  also  covers  many  small  areas  over  which  trees 
have  not  become  dominant.  Where  the  habitat  has  not  been  disturbed 
by  man  the  thick  tangle  of  smaller  shrubs,  thorns,  and  vines  makes 
excellent  refuges  for  birds  and  mammals. 

The  cottonwood-willow  association  is  made  up  of  a  great  number  of 
species.  A  large  number  of  these  are  closely  restricted  to  the  cotton- 
wood-willow  habitat.  A  few  species  which  reach  their  greatest 
abundance  in  the  cottonwood-willow  association  are  found  in  lesser 
abundance  in  the  bunchgrass  association.  Other  species  of  greatest 
abundance  in  the  bunchgrass  association  are  sparingly  represented 
in  the  cottonwood-willow  association.  Several  species  of  birds  nest  or 
obtain  shelter  in  the  cottonwood-willow  habitat  but  forage  out  into 
the  adjacent  bunchgrass. 

Sixty  traps  set  in  the  timber  and  brush  along  the  Touchet  River 
two  miles  east  of  Prescott  caught  on  July  2,  1914,  2  Reithrodontomys 
megalotis  nigrescens,  8  Peromyscus  maniculatus  gambelii,  and  5  Micro- 
tus  nanus  canescens. 


312 


University  of  California  Publications  in  Zoology        [V°L.  16 


WATEE-MAKGIN  HABITAT  AND  ASSOCIATION   (PRAIRIE  AREA) 

Exclusive : 

Chrysemys  bellii.  Actitis  macularius — summer. 

Grus  mexicana — summer. 

Major : 

Rana  pipiens  brachycephala. 
Thamnophis  elegans. 
Anas  platyrhynchos — winter. 
Nettion  carolinense — winter. 
Ardea  herodias  treganzai — resident. 
Oxyechus  vociferus  vociferus — 
resident. 

Minor : 

Spatula  clypeata — migrant. 
Streptoceryle  alcyon  caurina — 
resident. 

Reported : 

Bascanion  constrictor  vetustum. 


Agelaius  phoeniceus  neutralis — 

summer. 

Procyon  psora  pacifica. 
Mustela  vison  energumenos. 
Ondatra  zibethica  osoyoosensis. 
Castor  canadensis  canadensis. 


Riparia  riparia — summer. 
Mephitis  occidentalis  major. 
Peromyscus  maniculatus  gambelii. 


Along  the  smaller  streams  of  the  prairie  are  numerous  small  gravel 
and  dirt  bars.  These  usually  become  very  dry  in  summer  and  the 
grasses  and  herbs  growing  on  them  dry  up,  except  in  a  few  places  at 
the  level  of  the  water  or  along  the  rare  sloughs. 

Along  Snake  River  there  is  a  considerable  width  of  water-margin 
habitat,  which  is  annually  covered  during  the  spring  high  water. 
There  are  few  willows  or  shrubs  along  this  stream  and  the  water-margin 
habitat  is  broad  except  where  cliffs  reach  the  edge  of  the  water.  Near 
the  edge  of  the  water  plants  are  almost  absent,  only  a  few  herbs  being 
found.  On  the  higher  level  of  the  beach  there  is  considerable  drift- 
wood and  the  ground  is  quite  sandy.  In  among  the  logs  the  plants  of 
the  bunchgrass  habitat  appear  and  so  also  do  a  number  of  weeds. 

The  water-margin  association  of  the  prairie  is  chiefly  made  up  of 
species  which  feed  along  the  shores  of  the  streams.  Kingfishers  and 
bank  swallows  are  known  to  nest  in  holes  in  the  soft  dirt  banks. 
Robins  gather  mud  for  plastering  their  nests  from  along  the  shore. 
A  number  of  species  from  both  cottonwood-willow  and  bunchgrass 
associations  probably  come  to  the  water's  edge  to  drink. 

Twenty  traps  set  among  rocks  and  driftwood  on  the  shore  of  Snake 
River  near  Lyon's  Ferry  caught  two  Peromyscus  maniculatus  gambelii 
on  June  25,  1914. 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          313 


AQUATIC  HABITAT  AND  ASSOCIATION  (PRAIRIE  AREA) 

Exclusive : 

Charitonetta  albeola — migrant. 
Major : 

Spatula  clypeata — migrant.  Streptoceryle  alcyon  caurina — 

resident. 

Minor  : 

Anas  platyrhynchos — winter.  Mustela  vison  energumenos. 

Nettion  carolinense — winter.  Ondatra  zibethica  osoyoo^enms. 

Pandion  haliaetus  carolinensis —  Castor  canadensis  canadensis. 

summer. 

Reported : 

Mergus  americanus — winter.  Fulica  americana — migrant. 

Mareca  americana — migrant. 

The  Snake  River  is  a  large  stream  with  a  rapid  current,  but  the 
other  streams  of  the  prairie  area  are  small.  These  smaller  streams 
have  usually  a  rapid  current,  and  quiet  pools  are  rare.  Lakes  are 
entirely  absent. 

The  kingfisher,  osprey,  and  mink  feed  in  the  river  habitat.  Of  the 
other  forms  observed  on  the  rivers  some  probably  feed  in  the  habitat 
while  others  rest  or  take  refuge  there. 


AERIAL  HABITAT  AND  ASSOCIATION  (PRAIRIE  AREA) 

Major : 

Petroehelidon  lunifrons  lunifrons —          Myotis  yumanensis  (?). 

summer.  Myotis  californicus  californicus. 

Riparia  riparia — summer.  Lasiurus  cinereus. 

Stelgidopteryx  serripennis — 


Minor  : 

Tyrannus  tyrannus — summer.  Empidonax  difficilis  difficilis — 

Tyrannus  verticalis — summer.  summer. 

Sayornis  sayus — summer.  Empidonax  trailli  trailli — summer. 

Nuttallornis  borealis — migrant.  Empidonax  wrighti — summer. 

Myiochanes  richardsoni  richard-  Stetophaga  ruticilla — summer, 
soni — summer. 

Reported : 

Chordeiles  virginianus  hesperis —  Tachycineta  thalassina  lepida — 

summer.  summer. 

The  aerial  association  is  represented  in  the  prairie  area  by  night- 
hawks,  flycatchers,  swallows,  and  bats.  Some  of  the  swallows  are 
present  at  certain  localities  in  immense  numbers. 


314 


University  of  California  Publications  in  Zoology        [VOL.  16 


.     BLUE  MOUNTAIN  FAUNAL  AREA  AND  FAUNA 

Habitats  and  Associations: 

Rocky-slope.  Lowland-fir. 


Yellow-pine. 

Buckbrush. 

Alpine-fir. 


Water-margin. 

Aquatic. 

Aerial. 


The  Blue  Mountain  area  is  characterized  by  the  dominance  of 
conifer  forests  of  several  kinds.  In  the  bottoms  of  the  canons  the 
forest  is  often  very  heavy  and  is  dominated  by  the  lowland  fir  (Abies 
grandis).  On  the  tops  of  the  higher  ridges  the  alpine  fir  (Abies  lasio- 
carpa)  is  the  dominant  type  of  tree.  On  the  lower  ridges  and  slopes 
and  in  the  valleys  of  the  more  arid  parts  of  the  mountains  the  open 
yellow-pine  type  of  forest  prevails.  It  has  been  very  difficult  to  de- 
termine the  relation  of  the  species  of  vertebrates  to  the  different  kinds 
of  conifer  forest.  There  seems  to  be  a  restriction  of  certain  species  to 
the  higher  ridges  and  of  others  to  the  canons,  but  no  species  seems  to 
be  clearly  limited  to  any  particular  type  of  forest.  It  will  require  a 
very  considerable  amount  of  further  study  before  the  distribution  of 
the  vertebrates  in  the  area  is  at  all  satisfactorily  known. 

The  Blue  Mountains  are  at  the  present  time  only  partially  covered 
by  forests.  Many  of  the  steep,  rocky  slopes  are  nearly  or  quite  bare  of 
timber.  Also,  much  of  the  region  has  been  burned  over,  destroying  the 
forests.  Following  the  fires,  or  in  some  cases  starting  on  bare  slopes 
where  probably  no  fire  has  been,  there  have  sprung  up  extensive 
growths  of  deciduous  brush.  This  brush  is  best  developed  near  the 
summits  of  the  ridges,  but  extends  down  the  slopes  for  considerable 
distances,  reaching  the  bottoms  of  the  canons  at  the  heads  of  a  number 
of  streams. 

SPECIES  CHARACTERISTIC  OP  THE  BLUE  MOUNTAIN  FAUNA 


Rana  pretiosa. 
Charina  bottae. 

Dendrapagus  obscurus  richardsoni. 
Picoides  americanus  dorsalis. 
Sphyrapicus  thyroideus. 
Phloeotomus  pileatus  picinus. 
Stellula  calliope. 
Empidonax  hammondi. 
Perisoreus  canadensis  capitalis. 
Nucifraga  columbiana. 
Junco  hyemalis  shufeldti. 
Dendroica  townsendi. 


Cinclus  mexicana  unicolor. 
Nannus  hiemalis  pacificus. 
Sitta  pygmaea  pygmaea. 
Penthestes  gambeli  gambeli. 
Eegulus  satrapa  olivaceus. 
Myadestes  townsendi. 
Neosorex  navigator  navigator. 
Myotis  longicrus. 
Ursus  altifrontalis. 
Vulpes  macrourus. 
Martes  sp. 
Mustela  cicognanii  lepta. 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          315 


Evotomys  gapperi  saturatus. 
Microtus  mordax  mordax. 
Thomomys  fuscus  fuscus. 
Zapus  princeps  oregonus. 
Callospermophilus  chrysodeirus 


chrysodeirus. 

Sciurus  hudsonicus  richardsonii. 
Lepus  bairdii  bairdii. 
Odocoileus  hemionus  hemionus. 


The  Blue  Mountain  fauna  contains  a  number  of  species  which  in 
southeastern  Washington  are  not  found  in  the  other  faunal  areas. 
Many  of  these  species  are  characteristic  of  conifer  forests  elsewhere. 

ROCKY-SLOPE  HABITAT  AND  ASSOCIATIONS  (BLUE  MOUNTAIN~AREA) 

Exclusive : 

Salpinctes  obsoletus  obsoletus — summer. 
Major : 

Peromyscus  maniculatus  gambelii.  Callospermophilus  chrysodeirus 

chrysodeirus. 
Minor  : 

Falco  sparverius  sparverius — 

summer. 
Thomomys  fuscus  fuscus. 


Citellus  columbianus  columbianus. 
Eutamias  amoenus  amoenus. 


Outcroppings  of  basaltic  rock  are  common  in  the  Blue  Mountains. 
Besides  the  numerous  small  rock  bluffs  there  are  many  rocky  slopes 
covered  by  broken  pieces  of  rock  of  various  sizes.  These  rocky  slopes 
often  cover  large  areas.  On  exposed  slopes  grasses  and  small  shrubs 
such  as  ninebark  (Opulaster  pauciflorus)  grow  among  the  rocks  and 
in  many  places  there  are  scattered  yellow-pine  trees  (Pinus  ponde- 
rosa).  The  habitat  in  such  places  often  gradually  gives  way  to  the 
yellow-pine  forest  habitat. 

Seventy-eight  traps  set  on  a  rocky  slope  near  Hompeg  Falls  cap- 
tured 23  Peromyscus  maniculatus  gambelii  and  2  Eutamias  amoenus 
amoenus  on  July  24,  1914. 

YELLOW-PINE  HABITAT  AND  ASSOCIATION  (BLUE  MOUNTAIN  AREA) 


Minor : 

Dendrapagus  obscurus  richardsoni — 

resident. 
Falco  sparverius  sparverius — 

summer. 

Sphyrapicus  thyroideus — summer. 
Empidonax  wrighti — summer. 
Corvus  brachyrhynchos  hesperis — 

summer. 
Spizella  passerina  arizonae — 

summer. 

Junco  hyemalis  shufeldti — summer. 
Reported : 

Sialia  mexicana  occidentalis — summer. 


Piranga  ludoviciana — summer. 
Penthestes  gambeli  gambeli — 

summer. 
Planesticus  migratorius  propin- 

quus — summer. 
Sialia  currucoides — summer. 
Peromyscus  maniculatus  gambelii. 
Thomomys  fuscus  fuscus. 
Eutamias  amoenus  amoenus. 
Callospermophilus  chrysodeirus 

chrysodeirus. 


316  University  of  California  Publications  in  Zoology        [VOL.  16 

Over  the  foothills  and  exposed,  lower  slopes  of  the  Blue  Mountains 
the  yellow  pine  (Pinus  ponder osa)  forms  the  dominant  forest.  This 
species  seems  to  be  able  to  endure  much  drier  conditions  than  any  of 
the  other  conifers.  It  is  limited  in  vertical  range  and  is  not  found  on 
the  higher  parts  of  the  Blue  Mountains.  As  found  near  Hompeg 
Falls  yellow  pine  is  in  many  places  associated  with  Douglas  spruce 
(Pseudotsuga  taxi  folia).  The  trees  in  this  forest  usually  grow  rather 
far  apart.  On  the  exposed  slopes  there  is  little  underbrush,  but  the 
ground  is  stony  or  covered  by  grasses  and  prairie  plants.  On  some- 
what sheltered  slopes  a  considerable  amount  of  underbrush  may  be 
developed.  In  this  the  ninebark  (Opulaster  pauciflorus)  is  most 
abundant  and  in  places  on  eastern  slopes  forms  a  thick  covering  to 
the  ground.  Other  shrubs  such  as  service-berry  (Amelanchier),  cur- 
rants (Rubus),  spirea  (Spirea),  willows  (Salix),  and  alders  (Alnus) 
occur  also.  On  the  higher  slopes  the  forest  is  heavier  and  Douglas 
spruce  tends  to  become  dominant. 

No  trapping  was  done  in  the  yellow-pine  habitat  and  only  inci- 
dental observations  were  made  in  this  type  of  forest.  The  animal  in- 
habitants are  surely  much  more  numerous  than  indicated  in  the  above 
list. 

BUCKBEUSH  HABITAT  AND  ASSOCIATION  (BLUE  MOUNTAIN  AEEA) 

Exclusive : 

Taxidea  taxus  neglecta. 

Major : 

Buteo  borealis  calurus — summer.  Zapus  princeps  oregonus. 

Spizella  passerina  arizonae —  Eutamias  amoenus  amoenus. 

summer.  Lepus  bairdii  bairdii. 

Junco  hyemalis  shufeldti — summer.  Odocoileus  hemionus  hemionus. 
Thomomys  fuscus  fuscus. 

Minor : 

Colaptes  cafer  collaris — summer.  Citellus  columbianus  columbianus. 

Peromyscus  maniculatus  gambelii.  Callospermophilus  chrysodeirus 

Evotomys  gapperi  saturatus.  chrysodeirus. 

Microtus  mordax  mordax. 

Reported : 

Vulpes  macrourus.  Lynx  sp. 

The  principal  plant  of  the  buckbrush  habitat  is  the  buckbrush 
(Ceanothus  velutinus).  Associated  with  this  are  often  willows 
(Salix),  alders  (Alnus),  and  a  number  of  other  shrubs.  Near  the  tops 
of  the  ridges  stunted  aspens  (Populus  tremuloides)  sometimes  appear. 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          317 

The  brush  often  grows  so  thickly  that  it  is  very  difficult  to  force  one's 
way  through  it,  but  it  is  likely  to  be  in  clumps  and  there  are  many 
open  places.  Rock  outcroppings  are  numerous.  The  height  of  the 
brush  is  usually  from  four  to  eight  feet,  but  around  springs  or 
damp  places  it  may  grow  much  higher.  There  is  much  down  and 
partly  burned  timber  in  some  parts  of  the  habitat,  and  an  occasional 
stump  or  tree  has  survived  the  fires.  Young  conifers  are  springing  up 
in  places  and  of  these  the  lodgepole  pine  (Pinus  murrayana^ _is  most 
numerous  on  the  ridges.  On  the  lower  slopes  the  buckbrush  habitat 
overlaps  in  some  places  the  range  of  the  yellow  pine  (Pinus  ponder  osa] 
and  isolated  old  yellow  pine  trees  may  often  be  found  growing  in 
among  the  shrubs. 

Trapping  on  August  3  and  August  9,  1914,  in  buckbrush  on  the 
ridge  near  Twin  Buttes  R.S.  produced  1  Peromyscus  maniculatus  gam- 
belii,  1  Evotomys  gapperi  saturatus,  6  Zapus  princeps  or  eg  onus,  and  2 
Eutamias  amoenus  amoenus  from  a  total  of  50  trap-nights. 


ALPINE-FIE  HABITAT  AND  ASSOCIATION  (BLUE  MOUNTAIN  AREA) 

Exclusive  : 

Dryobates  villosus  orius— summer.  Sitta  canadensis — summer. 

Nucifraga  eolumbiana — resident. 


Major : 

Colaptes  cafer  collaris — summer. 
Spinus  pinus  pinus — summer. 
Penthestes  gambeli  gambeli — 


Sialia  currucoides — summer. 
Evotomys  gapperi  saturatus. 


Minor : 

Dendrapagus  obscurus  richard- 

soni — resident. 

Buteo  borealis  calurus — summer. 
Falco  sparverius  sparverius — 

summer. 

Sphyrapicus  thyroideus — summer. 
Cyanocitta  stelleri  annectens — 

resident. 
Spizella  passerina  arizonae — 

summer. 

Junco  hyemalis  shufeldti — summer. 
Nannus  hiemalis  pacificus — summer. 
Regulus  satrapa  olivaceus — summer. 

Reported  : 

Picoides  arcticus. 
Picoides  americanus  dorsalis — 
summer. 


Myadestes  townsendi — summer. 
Planesticus  migratorius  propin- 

quus — summer. 

Peromyscus  maniculatus  gambelii. 
Microtus  mordax  mordax. 
Thomomys  fuscus  fuscus. 
Zapus  princeps  oregonus. 
Eutamias  amoenus  amoenus. 
Callospermophilus  chrysodeirus 

chrysodeirus. 

Sciurus  hudsonicus  richardsonii. 
Lepus  bairdii  bairdii. 
Odocoileus  hemionus  hemionus. 


Passerella  iliaca  schistacea — 

summer. 

Ursus  altifrontalis. 
Lynx  sp. 


318  University  of  California  Publications  in  Zoology        [VOL-  16 

The  alpine  fir  (Abies  lasiocarpa)  forms  extensive  forests  on  the 
higher  ridges  of  the  Blue  Mountains.  It  does  not  grow  in  the  canons 
or  on  the  lower  slopes,  so  it  may  be  considered  the  characteristic  tree 
of  the  ridge  forest.  Alpine-fir  forest  is  abundantly  developed  in  many 
of  the  sheltered  coves  near  the  tops  of  the  ridges.  In  these  coves  the 
ground  is  fairly  moist.  The  forest  developed  is  not  dense,  and  there  is 
plenty  of  room  to  walk  between  the  trees.  The  size  reached  by  the 
alpine  firs  is  not  very  large  and  few  of  the  trunks  would  measure  over 
fifteen  inches  in  diameter.  Some  lodgepole  pines  (Finns  murrayana) 
are  often  mixed  with  the  alpine  firs  and  in  places  where  new  growth 
is  springing  up  lodgepole  may  be  the  dominant  tree.  There  is  usually 
very  little  undergrowth  under  the  alpine-fir  forest,  there  being  com- 
monly only  a  few  very  low  shrubs.  On  the  ridges  alpine  fir  is  less 
common  and  the  forest  is  more  open  and  commonly  includes  many 
Douglas  spruces  (Pseudotsuga  taxifolia).  The  ground  here  is  drier 
than  in  the  coves  and  there  is  much  exposure  to  sun  and  wind.  The 
trees  occur  singly  or  in  small  groups.  Douglas  spruce  is  the  dominant 
tree  in  the  most  exposed  places. 

In  many  of  the  coves  and  on  protected  slopes  near  the  tops  of  the 
ridges  forests  of  western  larch  (Larix  Occident alis)  are  developed. 
This  may  be  developed  as  a  pure  forest  or  may  be  mixed  with  alpine 
firs  or  Douglas  spruces.  The  pure  larch  forest  is  very  open  and  under- 
brush is  scanty  and  the  forest  floor  may  be  covered  entirely  by  grasses. 
The  trees  reach  a  good  size  with  trunks  several  feet  in  diameter.  On 
the  damper  slopes  Douglas  spruce  usually  dominates  over  the  larch 
and  a  rather  dense  forest  is  developed.  This  contains  much  more 
underbrush  and  the  ground  is  commonly  quite  damp.  In  such  places 
Engelmann  spruces  (Picea  Engelmanni)  may  occur  commonly. 

Although  the  alpine-fir  association  is  made  up  of  a  considerable 
number  of  forms  very  few  are  restricted  to  the  association.  Neither 
do  very  many  reach  their  greatest  abundance  in  this  habitat. 

Trapping  in  forest  habitats  on  the  ridges  near  Twin  Buttes  R.S. 
on  July  28,  July  29,  and  August  3,  1914,  produced  8  Peromyscus 
maniculatus  gambelii,  1  Microtus  mordax  mordax,  4  Evotomys  gap- 
peri  saturatus,  and  1  Eutamias  amoenus  amoenus  from  a  total  of  181 
trap-nights. 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          319 


LOWLAND-FIR  HABITAT  AND  ASSOCIATION  (BLUE  MOUNTAIN  AREA) 

Exclusive  : 

Phloeotomus  pileatus  picinus —  Penthestes  atricapillus  septentrion- 

summer. 
Melospiza  melodia  merrilli — 


alls — summer. 


Major : 

Dendrapagus  obscurus  richardsoni 

— resident. 
Falco  sparverius  sparverius — 

summer. 

Sphyrapicus  thyroideus — summer. 
Empidonax  difficilis  difficilis — 

summer. 

Empidonax  hammondi — summer. 
Empidonax  wrighti — summer. 
Nannus  hiemalis  pacificus — summer. 


Minor : 

Thamnophis  elegans. 
Colaptes  cafer  collaris — summer. 
Spinus  pinus  pinus — summer. 
Spizella  passerella  arizonae — 

summer. 

Junco  hyemalis  shufeldti — summer. 
Peromyscus  maniculatus  gambelii. 

Reported : 

Bufo  columbiensis. 

Charina  bottae. 

Bonasa  umbellus  togata — resident. 

Otus  asio  macfarlanei — summer. 

Stellula  calliope — summer. 

Passerella  iliaca  schistacea — 

summer. 

Dendroica  townsendi — summer. 
Oporornis  tolmiei— summer. 


Cyanocitta  stelleri 

resident. 
Corvus  brachyrhynchos  hesperis 

— summer. 

Piranga  ludoviciana — summer. 
Regulus  satrapa  olivaeeus — summer. 
Myadestes  townsendi — summer 
Planesticus  migratorius  propin- 

•quus — summer. 

Citellus  columbianus  columbianus. 
Sciurus  hudsonicus  richardsonii. 


Evotomys  gapperi  saturatus. 
Microtus  mordax  mordax. 
Thomomys  fuscus  fuscus. 
Eutamias  amoenus  amoenus. 
Castor  canadensis  canadensis. 
Lepus  bairdii  bairdii. 
Odocoileus  hemionus  hemionus. 


Certhia  familiaris  montana — 

summer. 
Penthestes  rufescens  rufescens — 

summer. 
Hylocichla  ustulata  swainsoni — 

summer. 

Sorex  vagrans  dobsoni. 
Martes  sp. 
Mustela  cicognanii  lepta. 


In  the  deeper  canons  of  the  Blue  Mountains  the  lowland  fir  (Abies 
grandis)  is  the  dominant  tree.  These  trees  reach  quite  a  large  size, 
trunks  estimated  at  over  four  feet  in  diameter  being  seen.  Near  Horn- 
peg  Falls  this  type  of  forest  is  well  developed.  However,  there  are 
many  open  places  washed  out  by  the  stream  or  due  to  the  action  of 
former  fires.  Associated  with  the  lowland  fir  are  Douglas  spruce 
(Pseudotsuga  taxifolia),  yellow  pine  (Pinus  ponderosa),  western  yew 


320  University  of  California  Publications  in  Zoology        [VOL.  16 

(Taxus  brevi folia),  western  larch  (Larix  Occident  alis),  cotton  wood 
(Populus  trichocarpa),  and  birches  (Betula  microphylla) .  A  few 
Englemann  spruces  (Picea  Englemanni)  occur  and  one  silver  pine 
(Pinus  monticola)  was  seen.  There  is  a  small  amount  of  underbrush, 
composed  chiefly  of  dwarf  maples  (Acer  glabrum)  and  alders  (Alnus 
sp.).  The  lowland -fir  type  of  forest  occurs  only  in  the  bottoms  of  deep 
canons  and  in  very  damp  places,  and  does  not  extend  up  on  the  moun- 
tain slopes. 

On  sheltered  lower  slopes  the  western  larch  and  Douglas  spruce 
make  up  the  larger  part  of  the  forest.  Sometimes  one  and  sometimes 
the  other  is  dominant.  The  larch  is  best  developed  in  damp  situations, 
while  the  Douglas  spruce  covers  drier  slopes.  Near  Hompeg  Falls  the 
larch-Douglas  spruce  forest  occupies  the  north  slopes  of  the  side 
ravines  which  branch  from  the  main  canon.  The  larch  largely  occupies 
the  bottoms  of  the  ravines,  while  the  Douglas  spruce  extends  further 
up  the  sides,  and  towards  the  tops  of  the  ridges  spreads  out  to  form 
a  more  extended  forest.  The  Douglas  spruce  forest  is  usually  fairly 
dense  in  this  situation  and  many  of  the  slopes  which  it  covers  are  very 
steep.  Under  the  heaviest  forest  of  this  kind  there  is  no  underbrush, 
but  the  ground  is  entirely  covered  by  dead  needles.  In  other  places 
the  forest  is  more  open  and  more  or  less  brush  occurs,  in  which  the 
alder  (Alnus  sp.)  is  the  most  abundant  type.  The  larch  forest  is  more 
open  and  usually  does  not  have  a  heavy  growth  of  underbrush. 

Along  Butte  Creek,  where  the  bottom  of  the  narrow  canon  has 
been  much  washed  over  by  the  stream,  much  of  the  lowland-fir  forest 
has  been  washed  out  and  is  replaced  in  patches  by  a  deciduous  forest 
which  is  notable  for  the  thickness  of  the  underbrush.  The  dominant 
trees  are  cottonwood  (Populus  trichocarpa)  and  the  willows  (Salix). 
The  brush  was  composed  principally  of  alder  (Alnus  sp.),  thorn 
(Crataegus  brevispina),  service-berry  (Amelanchier  florida),  wild 
cherry  (Primus  demissa),  red  osier  (Cornus  stolonifera) ,  dwarf  maple 
(Acer  glabrum),  and  snowberry  (Symphoricarpus) . 

Traps  set  in  the  lowland-fir  habitat  near  Hompeg  Falls  on  July  23, 
25,  and  26,  1914,  caught  1  Sorex  vagrans  dobsoni,  23  Peromyscus 
maniculatus  gambelii,  1  Evotomys  gapperi  saturatus,  and  2  Eutamias 
amoenus  amoenus  from  a  total  of  201  trap-nights. 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          321 

WATEE-MAKGIN  HABITAT  AND   ASSOCIATION 

(BLUE  MOUNTAIN  AEEA) 

Exclusive  : 

Eana  pretiosa.  Neosorex  navigator  navigator. 

Major: 

Thamnophis  elegans.  Castor  canadensis  canadensis. 

Microtus  mordax  mordax. 

Minor: 

Cinclus  mexicana  unicolor —  Evotomys  gapperi  saturatus. 

summer. 

Reported  : 

Telmatodytes  palustris  plesius — summer. 

Along  the  streams  of  the  Blue  Mountain  gravel  or  mud  bars  are 
rare  and  the  forest  often  reaches  the  edge  of  the  stream  and  partially 
overhangs  the  water.  Springs  are  quite  numerous  in  the  bottoms  of 
the  canons  and  these  often  produce  moist  areas  which  make  little 
swamps.  Such  swamps  are  often  shaded  by  the  heavy  lowland  fir 
trees  and  have  only  a  low  growth  of  vegetation,  but  in  more  open  spots 
grasses  and  horsetails  as  well  as  smaller  herbs  make  a  luxuriant  growth. 
Swamps  also  occur  along  small  sloughs  diverted  from  the  main  streams 
or  about  ponds  caused  by  the  damming  of  some  stream  by  a  beaver  dam 
or  by  the  natural  accumulation  of  drift. 


AQUATIC  HABITAT  AND  ASSOCIATION  (BLUE  MOUNTAIN  AEEA) 

Major  : 

Streptoceryle  alcyon  caurina —  Cinclus  mexicanus  unicolor — 

summer.  summer. 

Minor  : 

Castor  canadensis  canadensis. 


The  streams  in  the  Blue  Mountains  are  all  small  and  have  swift 
currents.  There  are  many  rapids  and  low  falls  and  the  pools  are 
small.  There  is  very  little  extent  of  open  water  free  from  the  margins 
of  the  streams. 


322  University  of  California  Publications  in  Zoology        [VOL.  16 

AERIAL  HABITAT  AND  ASSOCIATION  (BLUE  MOUNTAIN  AREA) 

Minor : 

Empidonax  difficilis  difficilis —  Empidonax  hammoiidi — summer, 

summer.  Empidonax  wrighti — summer. 

Reported : 

Chordeiles  virginianus  hesperis —  Myotis  longicrus. 

summer. 

Flycatchers,  nighthawks,  and  bats  make  up  the  members  of  the 
aerial  association  in  the  Blue  Mountains.  Swallows  were  seen  flying 
over  some  of  the  ridges  but  the  species  was  not  determined. 


CLIMATE 

Climatological  records  have  been  taken  for  a  number  of  years  in 
southeastern  Washington  by  Weather  Bureau  Stations.  The  accumu- 
lated data  have  been  kindly  furnished  by  the  United  States  Weather 
Bureau.  A  summary  of  this  is  presented  in  Table  1.  The  data  is  most 
complete  for  the  prairie  area  and  least  so  for  the  Blue  Mountains.  The 
stations  are  usually  located  in  towns  and  so  the  records  do  not  indicate 
the  conditions  in  any  particular  habitat,  but  they  do  give  a  basis  for 
comparing  the  climatic  conditions  in  different  faunal  areas. 

Table  2,  which  gives  the  climatological  data  for  each  month  at 
Walla  Walla,  is  presented  to  illustrate  the  weather  conditions  in  the 
region  at  the  different  seasons.  In  southeastern  Washington  the  pre- 
cipitation is  unequally  distributed  throughout  the  year,  being  greatest 
in  winter,  while  in  summer  very  little  rain  falls.  The  summers  are 
very  hot,  and  the  winters  moderately  cool  with  occasional  very  cold 
periods  of  short  duration.  There  is  a  considerable  daily  range  of 
temperature  and  even  in  the  hottest  weather  the  nights  are  cool.  The 
humidity  of  the  air  is  very  low  in  summer,  but  is  higher  in  winter. 
There  is  an  abundance  of  sunlight  in  summer,  while  in  winter  the 
light  is  much  weaker.  Winds  are  quite  common  and  especially  in 
spring  may  be  very  strong.  Their  usual  direction  is  from  the  south- 
west. A  peculiar  wind  which  deserves  notice  is  the  "chinook."  This 
is  a  dry,  warm  wind  from  the  southwest  which  may  start  at  any  time 
of  the  day  or  night  in  winter.  It  rapidly  melts  the  snow  and  dries  the 
surface  of  the  ground.  In  consequence,  snow  seldom  lies  for  any 
length  of  time  upon  the  ground,  except  in  the  Blue  Mountains. 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          323 


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1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          325 

The  growing  season  of  southeastern  Washington  is  comparatively 
long.  At  Prescott  it  is  often  possible  to  plant  the  seeds  of  hardy  vege- 
tables in  the  open  ground  in  the  first  week  of  March  or  earlier.  The 
frostless  season  is  also  comparatively  long  (Table  3),  although  irregu- 
lar frosts  late  in  spring  often  do  considerable  damage  to  fruit  and 
garden  crops. 

TABLE  III 
FROST  DATA 

Columbia  Basin  sagebrush  area  Columbia  Basin  prairie  area 

Station —  Kennewick  Touchet     Walla  Walla  Dayton 

Length  of  record  7  5  23  5 

Average   date   first   frost   in 

Autumn   Oct.    15  Sept.  13     Nov.     3  Oct.    13 

Average     date    last    killing 

frost  in  Spring Apr.    28  Apr.    23     Apr.      1  Apr.   25 

Earliest  date  of  killing  frost 

in  Autumn    Sept.  25  Aug.   25     Sept.  28  Sept.  23 

Latest  date  of  killing  frost 

in  Spring  May   25  May     8     May     3  May     8 

Average  season  between 

frosts— days  170  143  216  171 

Frostless  season — days 123  109  148  138 


No  records  of  humidity  are  available  from  the  various  habitats  of 
Walla  Walla  and  Columbia  counties,  but  in  Whitman  County,  Wash- 
ington, and  in  the  Thatuna  Hills  of  Idaho,  Weaver  (1914)  has  ob- 
tained records  of  the  rate  of  evaporation  as  determined  by  a  porous- 
cup  atmometer  during  the  summer.  These  records  show  that  the  rate 
of  evaporation  is  highest  in  the  rocky-slope  habitat  and  that  in  the 
other  habitats  it  decreases  in  the  following  order:  prairie,  S.W.  ex- 
posure ;  prairie,  N.E.  exposure ;  yellow  pine ;  fir-tamarack ;  cedar.  The 
sagebrush  habitat  was  not  included  in  these  observations. 

The  prominent  features  of  the  climate  of  the  sagebrush  area  is  the 
small  annual  precipitation  and  the  high  temperature  of  summer.  No 
records  of  wind  velocities  are  available  from  these  stations,  but  it  is 
known  that  the  area  is  subject  to  strong  winds  which  act  powerfully  to 
drift  the  sandy  soil. 

The  prairie  area  as  a  whole  shows  a  lower  average  temperature  and 
particularly  a  lower  temperature  in  the  summer  months  than  is  found 
in  the  typical  part  of  the  sagebrush  area  as  recorded  at  Kennewick. 
Also  the  prairie  area  shows  a  greater  rainfall  and  this  rainfall  is 
greater  the  more  closely  the  Blue  Mountains  are  approached.  It  may 


326  University  of  California  Publications  in  Zoology        [  V°L-  16 

be  said  also  that  the  winds  of  the  prairie  area  are  probably  less  strong 
than  those  of  the  sagebrush  area. 

Climatological  data  from  the  Blue  Mountains  is  very  scanty  and 
consists  only  of  records  of  precipitation  at  two  stations  on  the  lower 
ridges.  It  is  evident  that  the  precipitation  is  very  much  higher  on 
these  ridges  than  in  the  prairie  area.  The  temperature  of  the  area  is 
considerably  lower  than  in  the  adjacent  areas,  but  no  definite  records 
could  be  obtained.  There  is  a  much  greater  snowfall  in  the  mountains 
than  in  the  lower  country  and  the  snow  lies  on  the  mountains  all  winter 
and  often  until  late  in  the  spring. 


INFLUENCE  OF  ARTIFICIAL  CONDITIONS 

The  animal  habitats  of  southeastern  Washington  have  been  greatly 
altered  by  the  work  of  man.  Farming  is  extensively  carried  on  and  in 
the  prairie  area  a  very  large  percentage  of  the  land  is  under  cultiva- 
tion. Irrigation  is  also  practiced  in  the  valleys  of  both  the  prairie  and 
sagebrush  areas.  All  of  the  land  not  under  direct  cultivation  has  been 
heavily  grazed  by  cattle  and  stock.  Part  of  the  timber  along  the 
streams  has  been  cut  down  and  much  of  the  brush  has  been  cleared 
away.  Houses  have  been  built  and  shade  trees  planted  in  places 
where  formerly  no  trees  grew.  In  the  Blue  Mountains  there  have 
been  many  destructive  forest  fires  and  much  timber  has  been  cut.  In 
the  region  it  is  now  difficult  to  find  an  area  of  any  size  which  shows 
the  primitive  conditions  in  completeness. 

These  changes  in  the  environment  have  caused  great  changes  in  the 
abundance  of  the  different  species  of  vertebrates.  Some  species  are 
greatly  reduced  in  numbers  or  have  been  exterminated  in  the  region ; 
others  have  held  their  own  or  have  increased  to  some  extent.  The 
species  of  the  open  fields  have  probably  suffered  most  by  the  occupa- 
tion of  the  region  by  man.  Extensive  hunting  has  operated  to  reduce 
in  number  or  exterminate  some  of  the  game  animals.  On  the  other 
hand,  a  few  game  species  have  been  intentionally  introduced  by  man, 
and  a  few  obnoxious  species  have  been  unintentionally  introduced. 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          327 


COMPARISON  WITH  OTHER  SCHEMES  OF 
ECOLOGICAL  DISTRIBUTION 

The  vertebrate  associations  as  here  recognized  cannot  be  compared 
directly  with  other  schemes  of  vertebrate  associations,  because  the 
local  distribution  of  no  other  region  of  similar  climatic  conditions  has 
been  studied  by  the  associational  method.  However,  a  comparison  can 
be  made  with  several  schemes  of  ecologic  distribution  used  in  other 
regions. 

Weaver  (1914)  has  studied  the  plant  associations  found  in  Whit- 
man County,  Washington,  and  the  Thatuna  Hills  of  adjacent  Idaho, 
where  the  vegetational  features  are  somewhat  similar  to  those  of  the 
region  we  are  studying.  He  recognizes  the  following  plant  associa- 
tions: bunchgrass-rimrock  association,  prairie  association,  yellow  pine 
association,  fir-tamrack  association,  and  cedar  association.  The  bunch- 
grass-rimrock association  corresponds  to  the  rocky-slope  association 
of  the  prairie  area  as  used  in  this  paper.  The  prairie  association  is 
the  same  as  our  bunchgrass  association. 

Gates  (1911,  pp.  9-11)  has  included  flycatchers  and  swallows  in 
the  aquatic  association  because  they  capture  insects  in  the  air  over 
the  water.  However,  such  forms  cannot  be  considered  to  be  aquatic  in 
any  sense  of  the  term  and  we  have  therefore  placed  them  in  the  aerial 
association. 

Shelf ord  (1913,  p.  262)  in  the  Chicago  area  has  recognized  a  dis- 
tinct animal  community  in  the  narrow  border  of  shrubs  and  weeds 
occurring  between  the  prairie  and  the  forest  proper.  This  forest- 
margin  community  is  very  distinct  in  many  regions,  but  it  has  been 
thought  undesirable  to  recognize  it  as  an  association  between  the  willow 
associations  and  the  bunchgrass  or  sagebrush  associations  of  south- 
eastern Washington.  The  willow  habitat  in  the  region  is  usually  nar- 
row and  is  often  rather  open.  It  resembles  in  these  respects  the  forest 
margin  rather  than  a  true  forest  habitat.  In  the  Blue  Mountains  the 
yellow  pine  forests  pass  over  into  the  prairie  usually  without  any  indi- 
cation of  a  marginal  habitat.  The  other  conifer  forests  of  the  area  are 
sometimes  bordered  by  an  extensive  growth  of  brush  and  this  has  been 
called  the  buckbrush  habitat. 

Kennedy  (1914)  in  a  study  of  the  birds  of  the  Yakima  Valley, 
Washington,  gives  separate  lists  of  the  birds  of  the  sagebrush  and  of 
those  found  along  the  streams.  No  attempt  is  made  to  distinguish 


328  University  of  California  Publications  in  Zoology        [VOL.  16 

those  of  the  timber  and  brush  from  those  of  the  stream  shore.  The 
Yakima  Valley  belongs  in  the  Columbia  Basin  sagebrush  faunal  area 
and  the  environmental  conditions  of  the  sagebrush  in  that  valley  seem 
to  be  very  similar  to  those  of  the  sagebrush  in  western  Walla  Walla 
County.  Of  the  species  of  birds  stated  to  be  characteristic  of  the  sage- 
brush of  the  Yakima  Valley  all  except  five,  Sayornis  say  us,  Otocoris 
alpestris  merrilli,  Pooecetes  gramineus  confinus,  Spizella  breiveri,  and 
Oreoscoptes  montanus,  have  been  reported  from  the  sagebrush  of 
western  Walla  Walla  County. 

The  roadside  association  recognized  by  Jackson  (1914,  pp.  23,  24) 
in  the  conifer  forests  of  Wisconsin  belongs  to  a  habitat  at  the  edge  of 
a  clearing  in  a  heavy  forest  and  seems  to  have  many  features  in  com- 
mon with  the  forest-margin  communities  recognized  in  other  regions. 
Such  an  association  might  be  recognizable  in  the  Blue  Mountains,  but 
roads  are  few  in  that  area  and  it  is  impossible  to  define  such  an  asso- 
ciation without  more  data  than  is  at  present  at  hand. 

Animal  habitats  are  sometimes  divided  into  strata.  Shelf ord  (1913, 
p.  165)  recognizes  five  strata  in  some  terrestial  habitats,  extending 
from  the  subterranean  stratum  to  the  tree  stratum.  No  attempt  has 
been  made  to  divide  the  habitats  of  southeastern  Washington  into 
strata,  although  various  strata  could  undoubtedly  be  distinguished. 

Much  has  been  made  of  the  succession  of  animal  species  due  to  the 
change  in  habitats  induced  or  correlated  with  plant  succession  ( Adams, 
1908).  In  southeastern  Washington  many  of  the  associations  and 
habitats  seem  to  have  reached  an  equilibrium  and  succession  is  not 
very  prominent.  In  the  sagebrush  and  prairie  areas  the  rocky-slope 
habitat  tends  to  change  to  the  sagebrush  or  bunchgrass  habitat.  Modi- 
fications which  occur  by  the  shiftings  of  the  stream  channels  produce 
changes  in  the  riparian  associations.  Floods  sometimes  wash  out  part 
of  the  willow  habitat  and  even  at  times  part  of  the  bunchgrass  or  sage- 
brush habitat.  Also,  the  willow  habitat  tends  to  invade  the  river  beds. 
At  every  shifting  of  the  stream  channel  there  are  changes  in  the 
extent  and  position  of  the  water-margin  habitat.  In  the  Blue  Moun- 
tains the  conditions  are  probably  less  stable  and  changes  in  habitats 
are  probably  in  more  active  progress.  Weaver  (1914)  has  suggested 
that  in  Whitman  County,  Washington,  and  in  the  adjacent  parts  of 
Idaho  the  succession  is  the  following  direction:  (1)  bunchgrass;  (2) 
yellow  pine;  (3)  Douglas  spruce  and  western  larch;  (4)  cedar.  Cedar 
does  not  occur  on  the  Blue  Mountains  as  a  distinct  habitat,  but  its 
place  is  probably  taken  by  the  alpine  fir. 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          329 

Associations  may,  for  ease  in  comparison,  be  grouped  in  either  of 
several  different  manners.  Grinnell  and  Swarth  (1913,  pp.  218-220) 
have  considered  two  kinds  of  associations,  major  and  minor.  Each 
major  association  is  made  up  of  one  or  more  minor  associations.  Major 
associations  recognized  in  the  San  Jacinto  area  of  California  are : 
chaparrel,  forest,  riparian,  rupestrine,  meadow  and  sand-flat.  It  is 
considered  that  a  given  major  association  may  occur  in  several  faunal 
areas  and  life-zones,  but  its  minor  divisions  are  much  more  restricted. 
The  associations  of  southeastern  Washington  recognized  in  this  paper 
would  belong  in  general  to  the  -class  of  major  associations  according 
to  this  classification,  for  no  attempt  has  been  made  in  most  cases  to 
work  out  the  finer  divisions  of  the  associations. 

Another  method  of  comparing  associations  is  to  group  them  into 
formations.  A  formation  is  stated  to  be  a  group  of  physiologically 
similar  associations  ( Shelf ord,  1913,  p.  38).  Formations  may  them- 
selves be  combined  into  still  larger  groups.  The  classification  of  the 
formations  of  the  world  is  still  in  its  preliminary  stages. 

Shelf  ord  (1911,  pp.  604,  605)  has  proposed  a  classification  of 
formations  with  Avhich  it  will  be  illuminating  to  compare  the  associa- 
tions of  southeastern  Washington.  The  conifer  forest  associations  of 
the  Blue  Mountains  must  be  referred  to  his  second  division,  forma- 
tions of  forests  with  narrow,  thick  leaves.  The  bunchgrass  associations 
belong  to  the  third  division,  formations  of  savannas  and  grasslands, 
and  to  the  subdivision  c,  cool  steppe  formations.  The  associations  of 
the  rivers  belong  to  division  seven,  formations  of  fresh  water.  The 
other  associations  are  harder  to  place  in  the  system. 


ZOOGEOGRAPHIC  POSITION  OF  SOUTHEASTERN 
WASHINGTON 

The  accompanying  table  (Table  4)  shows  the  general  relations  of 
the  vertebrate  faunas  of  southeastern  Washington  to  the  faunas  of 
adjacent  regions.  In  this  table  the  occurrences  in  the  adjacent  regions 
are  given  of  those  species  whose  ranges  are  well  known  and  which  have 
been  definitely  identified  from  Walla  Walla  County  or  Columbia 
County.  Of  the  birds  only  those  species  occurring  in  the  regions  in 
summer  and  which  are  presumably  breeding  are  included. 

The  Columbia  Basin  sagebrush  fauna  shows  in  this  comparison 
much  greater  affinity  to  the  fauna  of  the  Great  Basin  than  to  the 


330  University  of  California  Piiblications  in  Zoology        tv°L- 16 


No.  of 


TABLE  IV 

Occurring  in  adjacent  districts  Unique  in 


Occurring  in  S.E.  forms  Rocky  Cascade  Great  Puget      Columbia         Blue 

Washington  in :  considered  Mts.  Mts.  Basin  Sound         Basin             Mts. 

Sagebrush  fauna  only....  11  1083  3 

Prairie  fauna  only 29  20  4  23  15                1 

Blue  Mountain  fauna 

only  35  30  18  3  10-                                1 

In  both  sagebrush  and 

prairie  faunas  34  25  5  34  14 

In  both  prairie  and  Blue 

Mountain  faunas 17  14  8  7  7 

In  sagebrush,  prairie,  & 

Blue  Mt.  faunas....  16  12  6  15  7 


faunas  of  other  adjacent  regions.  It  must  be  placed  in  the  Great 
Basin  district  of  Allen  (1892,  p.  237).  As  the  Great  Basin  seems  to 
have  had  a  rather  stable  climate  and  a  continuous  sequence  of  forms 
for  a  long  period  of  geologic  time  it  is  allowable  to  suppose  that  many 
of  the  forms  of  the  Columbia  Basin  sagebrush  fauna  originated  in  the 
Great  Basin  and  migrated  into  southeastern  Washington  at  some 
period  later  than  the  Middle  Miocene. 

The  fauna  of  the  Blue  Mountains  is  most  closely  related  to  the 
fauna  of  the  Rocky  Mountains.  The  Blue  Mountain  area  must  be 
placed  in  a  subdivision  of  the  Canadian  subregion  of  Holarctica 
(Lydekker,  1896,  p.  360).  It  seems  logical  to  suppose  that  the  fauna 
of  this  area  has  been  derived  largely  from  the  North. 

The  fauna  of  the  Columbia  Basin  prairie  area  is  related  to  -the 
faunas  of  both  the  Rocky  Mountains  and  the  Great  Basin.  It  has 
seemingly  been  produced  largely  by  an  admixture  of  elements  from 
these  two  places. 

The  maintainance  of  the  distinctness  of  the  fauna  of  the  Columbia 
Basin  prairie  area  must  be  due  to  the  climatic  barriers  which  separate 
it  from  the  Columbia  Basin  sagebrush  fauna  and  from  the  Blue  Moun- 
tain fauna.  Differences  in  temperature  and  rainfall  and  perhaps 
other  factors  are  effective  in  separating  the  fauna  of  the  prairie  from 
that  of  the  Blue  Mountains.  The  difference  in  temperature  between 
the  prairie  area  and  the  sagebrush  area  in  southeastern  Washington 
is  not  marked  and  the  difference  in  rainfall  is  probably  the  chief  factor 
separating  the  faunas  of  the  two  places. 


1916]-       Dice:  Land  Vertebrates  of  Southeastern  Washington          331 


LIFE-ZONES  OF  SOUTHEASTERN  WASHINGTON 

The  sagebrush  region  about  Wallula  belongs  certainly  to  the  Upper 
Austral  life-zone.  It  is  placed  in  this  zone  on  the  basis  of  the  flora 
by  Piper  (1906,  p.  35).  Merriam  (1898,  p.  30)  states  that  a  part  of 
the  Upper  Austral  zone  in  Washington,  in  the  valleys  of  the  Snake 
and  Columbia  rivers,  has  so  hot  a  climate  that  it  might  almost  be 
placed  in  the  Lower  Austral  zone. 

The  Columbia  Basin  prairie  area  must  be  placed  in  the  Transition 
life-zone  although  it  contains  a  strong  Upper  Sonoran  element.  In 
the  area  there  are  no  species  which  have  not  elsewhere  been,  r-eported 
to  occur  in  zones  above  the  Upper  Sonoran.  Four  breeding  species, 
Mustela  arizonensis,  Citellus  columbianus  columbianus,  Passerella 
iliaca  scliistacea,  and  Oporornis  tolmiei,  are  characteristic  of  the 
Transition  or  higher  life-zones.  Further,  the  area  has  been  placed  in 
the  Transition  life-zone  by  Piper  (1906,  p.  48)  on  the  basis  of  the 
flora. 

In  the  Transition  life-zone  must  be  included  the  bunchgrass  hills 
south  of  Wallula  and  also  those  north  of  the  Walla  Walla  River  east 
of  Nine-mile.  Sagebrush  as  a  dominant  habitat  extends  up  the  Walla 
Walla  Valley  as  far  as  Touchet,  and  this  would  seem  to  mark  the 
eastern  limit  of  the  Upper  Austral  life-zone  in  the  region.  Piper 
(1906,  map)  extends  a  tongue  of  Upper  Austral  as  far  east  as  Walla 
Walla,  but  there  seems  no  justification  for  this,  for  the  plant  and  ani- 
mal associations  at  Walla  Walla,  so  far  as  can  be  judged  under  the 
present  altered  conditions,  are  essentially  the  same  as  in  the  bunch- 
grass  region  to  the  north  and  east.  Piper  also  places  the  canon  of 
Snake  River,  for  the  whole  of  the  distance  that  this  extends  through 
Washington,  in  the  Upper  Austral  life-zone.  However,  in  the  canon 
of  Snake  River  at  Lyon  's  Ferry  sagebrush  was  not  the  dominant  vege- 
tation and  the  characteristic  vertebrates  of  the  Upper  Austral  life- 
zone  found  at  Wallula  were  not  present. 

Temperature  records  of  the  kind  used  by  Merriam  (1894)  in  de- 
fining the  limits  of  the  life-zones  are  available  only  for  Walla  Walla 
(Bigelow,  1908,  p.  90).  At  Walla  Walla  daily  normal  temperatures 
of  43°  F.  and  above  occur  throughout  the  period  between  March  12 
and  November  16,  giving  an  average  growing  season  of  249  days.  The 
sum  of  the  daily  normal  temperatures  for  this  season  is  15352°  F. 
The  hottest  six  weeks  of  summer  at  Walla  Walla  are  the  last  three 
weeks  of  July  and  the  first  three  weeks  of  August.  The  average  tern- 


332  University  of  California  Publications  in  Zoology        [VOL.  16 

perature  of  this  period  is  74.8°  F.  On  the  basis  of  temperature  it 
would  be  necessary  to  place  Walla  Walla  and  the  Columbia  Basin 
prairie  area  about  midway  in  the  Upper  Austral  life-zone  (Merriam, 
1894,  p.  236),  but  the  fauna  indicates  closer  affinity  to  the  Transition 
life-zone.  In  this  matter  the  fauna  is  probably  a  better  criterion  than 
the  temperature,  because  the  life-zones  are  founded  primarily  on 
faunal  relationships. 

The  yellow-pine  areas  of  the  lower  parts  of  the  Blue  Mountains  in 
Columbia  County  make  up  the  timbered  division  of  the  Transition 
life-zone  as  recognized  by  Piper  (1906,  p.  35). 

The  part  of  the  Blue  Mountain  area  above  the  Transition  life-zone 
belongs  to  the  Boreal  region  of  Merriam.  Piper  (1906,  pp.  58,  60,  62) 
recognizes  Canadian,  Hudsonian,  and  Arctic  life-zones  in  the  flora  of 
these  mountains.  In  the  vertebrate  fauna  the  Arctic  life-zone  cannot 
be  distinguished  and  the  Canadian  and  Hudsonian  life-zones  are  very 
difficult  to  separate.  If  the  Hudsonian  life-zone  be  recognized  as  dis- 
tinct it  must  be  restricted  to  the  summits  of  the  ridges.  Here  is  found 
the  alpine  fir  (Abies  lasiocarpa),  a  characteristic  Hudsonian  tree 
(Piper,  1906,  p.  60).  The  vertebrate  species  found  on  these  higher 
ridges,  and  not  reported  from  lower  altitudes,  are  Zapus  princeps  ore- 
gonus,  Picoides  americanus  dorsalis,  and  Nucifraga  columbiana.  None 
of  these  species  can  be  considered  strictly  Hudsonian.  It  seems  best 
to  place  the  part  of  the  Blue  Mountain  area  above  the  Transition  life- 
zone  in  a  single  life-zone,  the  Boreal. 


COMPARISON  OF  THE  DIFFERENT  SYSTEMS  OF 
CONSIDERING  DISTRIBUTION 

The  facts  of  animal  and  plant  distribution  are  very  complex  and  it 
is  convenient  to  have  some  system  or  systems  of  arranging  these  facts 
so  that  they  can  be  considered  in  groups  rather  than  as  isolated  in- 
stances. Several  systems  are  now  in  use.  Each  of  these  emphasizes 
different  features  of  the  facts  of  distribution. 

THE  ZOOGEOGRAPHICAL  SYSTEM 

The  system  of  zoogeography  points  out  the  barriers  to  distribution, 
and  indicates  something  as  to  the  origin  of  the  faunas  of  different 
regions.  Because  different  species  of  animals  are  not  limited  by  the 
same  barriers,  they  do  not  all  fall  evenly  into  zoogeographical  divisions. 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          333 

Routes  of  migration  have  been  opened  and  closed  at  irregular  times, 
and  many  groups  have  become  differentiated  only  to  become  exter- 
minated. Climatic  barriers  are  hard  to  determine  and  different 
species  show  various  degrees  of  limitation  by  such  barriers.  It  is  a 
difficult  matter  to  divide  any  region  into  satisfactory  zoogeographical 
areas.  It  has  been  pointed  out  that  the  zoogeographical  divisions  of 
the  globe  are  different  for  each  group  of  animals  and  that  in  any  one 
group  these  divisions  indicate  roughly  the  length  of  time  the  different 
sections  of  the  group  have  been  separated  (Gadow,  1913,  p.~13~15). 

The  zoogeographical  method  has  many  limitations  and  it  is  unwise 
to  attempt  to  apply  it  too  closely.  Species  are  limited  in  distribution 
by  various  factors  or  complexes  of  factors.  The  zoogeographical 
divisions  are  founded  on  comparative  statistics  and  there  will  always 
be  exceptions.  In  some  cases  the  exceptions  will  almost  equal  the 
number  following  the  rule.  The  zoogeographical  divisions  are  more 
or  less  arbitrary  and  there  are  sure  to  be  many  places  of  uncertain 
position.  Still,  the  system  greatly  simplifies  the  consideration  of  the 
facts  of  distribution. 


THE  LIFE-ZONE  SYSTEM 

The  significance  of  the  life-zone  method  lies  in  its  indication  of 
climatic  barriers  on  the  continent  of  North  America,  and  the  origin  of 
the  faunas  of  the  several  life-zones.  Although  this  method  of  con- 
sidering distribution  has  come  into  rather  general  use,  it  has  a  number 
of  disadvantages  and  difficulties,  and  a  considerable  amount  of  criti- 
cism has  been  directed  at  the  system.  It  seems  advisable  therefore  to 
consider  its  history  and  some  of  the  objections  which  have  been  raised 
against  it. 

On  the  high  mountains  of  the  western  United  States  there  are  dif- 
ferent zones  of  vegetation  at  different  levels,  and  with  each  of  these 
vegetational  zones  there  are  associated  particular  species  of  animals. 
Merriam  (1890,  pp.  7-11)  found  seven  such  zones  of  life  on  San  Fran- 
cisco Mountain,  Arizona.  Beginning  at  the  top  he  gave  these  the 
names  of  Alpine  zone,  Subalpine  or  Timberline  zone,  (Central)  Hud- 
sonian  or  Spruce  zone,  (Central)  Canadian  or  Balsam  Fir  zone, 
Neutral  or  Pine  zone,  Pinon  zone,  and  the  Desert  Area.  He  showed 
that  some  species  in  the  fauna  and  flora  of  the  uppermost  four  of 
these  zones  were  characteristic  of  much  more  northern  regions.  On 
the  mountains  of  central  Idaho,  Merriam  (1891,  pp.  21-25)  distin- 


334  University  of  California  Publications  in  Zoology        [VOL-  16 

guished  six  life-zones  which  he  called  respectively  the  Arctic-alpine 
zone,  the  Subalpine  or  Timberline  zone,  the  (Central)  Hudsonian  or 
Spruce  zone,  the  (Central)  Canadian  or  Douglas  Fir  zone,  the  Neutral 
or  Transition  zone,  and  the  Upper  Sonoran  zone. 

The  faunal  divisions  of  eastern  North  America  generally  recognized 
by  students  of  distribution,  particularly  by  ornithologists,  at  the  time 
Merriam  began  his  work  on  correlation,  were  eight  in  number.  Pass- 
ing from  north  to  south  these  divisions  were  (1)  Arctic,  (2)  Hud- 
sonian, (3)  Canadian,  (4)  Alleghanian,  (5)  Carolinian,  (6)  Louis- 
ianian,  (7)  Floridian,  and  (8)  Antillean  (Merriam,  1890,  p.  18). 

Merriam  (1890,  p.  18)  was  much  impressed  with  the  similarities 
between  the  zones  of  the  higher  parts  of  San  Francisco  Mountain  and 
the  faunal  areas  of  northeastern  North  America  and  states  that 

in  many  instances,  the  zones  of  the  mountain  may  be  recognized  by  the  identical 
species  which  characterize  them  in  New  England  and  Canada.  In  short  it  was 
found  that  the  faunal  and  floral  zones  which  go  to  make  up  the  Boreal  province 
in  the  East  may  be  traced  in  a  northwesterly  direction  around  the  northern  end 
of  the  Plains  of  the  Saskatchewan,  and  then  south  along  the  sides  of  the  Rocky 
Mountains  even  to  this  isolated  peak  in  Arizona. 

Merriam  (1892,  p.  22)  later  extended  the  correlation  of  the  zones 
of  the  eastern  and  western  United  States  and  stated  "with  some  con- 
fidence" that  the  Transition  zone  of  the  mountains  of  the  West  is  the 
equivalent  of  the  Alleghenian  of  the  East  and  also  that  the  Upper 
Sonoran  is  the  equivalent  of  the  Carolinian,  and  the  Lower  Sonoran  of 
the  Austroriparian.  He  thought  that  these  life-zones  followed  "the 
lines  of  equal  temperature  during  the  season  of  reproduction,"  and 
based  the  correlation  mainly  on  that  factor. 

Since  that  time  members  of  the  United  States  Bureau  of  Biological 
Survey  and  others  have  extensively  followed  the  life-zone  method  in 
describing  distribution  in  North  America.  A  brief  statement  of  the 
birds  and  mammals  characteristic  of  each  life-zone  was  published  by 
Merriam  in  1898.  As  used  as  present,  the  Timberline  zone  originally 
recognized  by  Merriam  has  been  merged  into  the  Hudsonian  zone,  but 
no  other  important  modification  has  been  made. 

Each  of  the  life-zones  of  the  Sonoran  region  is  divisible  into  two 
or  more  faunal  areas  (Merriam,  1898,  pp.  20-49).  These  faunal 
divisions  are  based  upon  differences  in  the  atmospheric  humidity  in 
different  parts  of  the  same  life-zone  (Grinnell  and  Swarth,  1913,  p. 
217).  In  California  a  considerable  number  of  these  faunas  have  been 
distinguished  by  Grinnell  (1902,  p.  7). 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          335 

The  idea  that  temperature  is  the  fundamental  factor  in  limiting 
the  distribution  of  species  is  dominant  in  the  conception  of  life-zones, 
and  in  1894  Merriam  made  an  attempt  to  determine  the  temperature 
limits  of  each  life-zone.  By  running  various  isothermal  lines  he  de- 
termined that  the  northern  limits  of  the  life-zones  agreed  fairly  well 
in  having  the  same  total  quantity  of  heat.  The  total  quantity  of  heat 
is  the  sum  for  the  year  of  the  daily  mean  temperatures  above  6°  C. 
This  temperature  is  assumed  to  be  the  point  at  which  life  begins 
activity.  The  southern  limits  of  the  life-zones,  however,  did  not  agree 
with  the  isotherms  thus  determined,  but  did  approximately  agree  with 
isotherms  of  the  hottest  period  of  the  year. 

One  criticism  which  should  be  made  of  this  correlation  of  life-zones 
and  isotherms  is  that  the  northern  and  southern  limits  of  the  life-zones 
are  not  determined  by  the  same  temperature  criteria  and  that  there- 
fore in  some  places  the  life-zones  may  not  meet  each  other.  If  de- 
pendence be  placed  on  these  temperature  criteria  alone,  some  regions 
must  be  placed  in  two  life-zones  and  theoretically  some  in  none  at  all. 
Along  the  Pacific  Coast  in  particular  there  is  much  overlapping  of  the 
life-zones.  Merriam  (1894,  p.  233-235)  considers  that  in  that  region 
the  northern  forms  are  able  to  come  far  south  on  account  of  the  low 
temperature  of  the  summers,  while  the  southern  forms  are  able  to 
extend  their  ranges  far  to  the  north  on  account  of  the  long  growing 
season.  Thus  is  explained  the  great  overlapping  of  northern  and 
southern  forms  in  the  "Pacific  Coast  strip."  However,  it  cannot  be 
considered  proved  that  the  temperature  relations  established  by 
Merriam  are  the  particular  ones  which  determine  the  limits  of  dis- 
tribution of  any  species  of  animal. 

Another  criticism  of  Merriam 's  determination  of  life-zone  tem- 
peratures is  that  no  thorough  attempt  has  been  made  to  determine  if 
these  temperatures  actually  do  apply  to  all  parts  of  the  life-zones  as 
they  have  been  plotted  in  North  America.  Indeed,  the  temperatures 
of  some  parts  of  the  life-zones  in  the  West  were  obtained  by  applying 
temperature  data  obtained  in  the  corresponding  faunal  areas  of  the 
eastern  United  States  (Merriam,  1890,  pp.  31,  32).  There  are  some 
facts  which  seem  to  indicate  that  the  temperatures  determined  by 
Merriam  do  not  apply  in  parts  of  some  life-zones.  For  instance,  ac- 
cording to  temperature  Walla  Walla  and  the  Columbia  Basin  prairie 
area  of  southeastern  Washington  would  be  placed  well  within  the 
Upper  Austral  zone,  but  the  faunal  relationships  are  with  the  Transi- 
tion zone,  or  at  least  are  not  definitely  Upper  Austral. 


336  University  of  California  Publications  in  Zoology        [VOL. 16 

In  the  life-zone  system,  humidity  is  recognized  to  have  considerable 
influence  on  distribution,  but  is  held  to  be  always  subordinate  to  the 
influence  of  temperature.  However,  it  seems  that  either  of  several 
climatic  factors  may  be  of  importance  in  limiting  organic  distribution. 
An  extreme  variation  of  humidity,  or  probably  of  other  climatic  fac- 
tors besides  temperature,  may  form  a  positive  barrier  to  the  distribu- 
tion of  species.  All  the  climatic  factors  are  complexly  interrelated 
and  a  variation  of  any  factor  has  an  influence  on  the  effect  of  the 
others.  Different  organisms  are  adapted  to  different  climatic  com- 
plexes and  react  in  different  manners  to  different  factors  and  to  vary- 
ing degrees  of  the  same  factor.  Temperature  perhaps  often  is  the 
most  important  factor  in  limiting  distribution,  but  it  would  seem  to 
be  impossible  to  base  a  system  of  distribution  on  variations  in  any  one 
climatic  factor  without  obscuring  many  facts  of  prime  importance.  It 
has  not  yet  been  established  that  small  differences  of  temperature  of 
the  degree  supposedly  separating  some  of  the  life-zones  are  as  im- 
portant barriers  to  distribution  as  are  some  of  the  more  marked  dif- 
ferences due  to  variations  in  rainfall  and  humidity. 

The  zones  of  life  which  occur  in  any  given  locality  may  be  depend- 
ent in  part  on  temperature,  yet  there  are  other  factors  which  evidently 
have  a  very  strong  modifying  influence.  Differences  in  rainfall,  in 
the  humidity  of  the  air,  in  slope  exposure,  or  in  other  factors  may 
greatly  modify  the  position  of  zones.  It  may  be  that  differences  in 
some  of  those  factors,  other  than  temperature,  might  even  be  the 
principal  cause  in  the  production  of  certain  zones.  In  each  given  case 
it  is  probably  the  complex  of  climatic  factors  which  determines  the 
occurrence  of  the  zone  rather  than  the  action  of  one  factor  alone. 

Three  distinct  regions  of  life  may  be  recognized  in  North  America, 
the  Holarctic  (Boreal)  region,  the  Sonoran  (Austral)  region,  and  the 
Neotropical  region  (Tropical  zone)  (Lydekker,  1896,  frontispiece). 
The  limitation  of  many  characteristic  species  and  genera  to  each  of 
these  regions  is  probably  due  principally  to  the  action  of  temperature 
as  a  barrier.  In  the  Holarctic  region  of  North  America  three  trans- 
continental belts  of  life,  Arctic,  Hudsonian,  and  Canadian,  have  been 
recognized  by  nearly  all  students  of  geographical  distribution.  These 
belts  of  life  are  probably  also  determined  largely  by  the  effect  of 
temperature.  However,  there  is  much  more  difficulty  in  recognizing 
transcontinental  life  belts  within  the  Sonoran  region,  and  in  the  truly 
tropical  regions  zones  of  distribution  corresponding  to  isotherms  have 
not  been  recognized. 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          337 

Zones  of  life  are  clearly  evident  upon  many  mountains  and  in  many 
regions  which  are  not  mountainous.  The  life  of  the  uppermost  of  some 
of  these  mountain  zones  is  evidently  related  to  the  life  of  more  north- 
ern regions.  However,  the  life  of  the  mountain  zones  is  never  identical 
with  the  life  of  any  particular  northern  transcontinental  belt.  Neither 
do  the  zones  found  on  mountains  in  different  parts  of  the  United  States 
exactly  correspond.  Grinnell  finds  the  Canadian  and  Hudsonian  life- 
zones  in  California  to  be  far  less  distinct  than  the  other  life^zones  in 
the  state.  Also  in  California  those  two  life-zones  are  much  less  dis- 
tinct than  they  are  in  the  northern  part  of  the  continent.  In  the  Blue 
Mountains  of  Washington  it  is  almost  impossible  to  separate  the  Hud- 
sonian life-zone  from  the  Canadian.  In  the  Pine  Forest  Mountains  of 
Nevada,  Taylor  (1912,  p.  339)  recognizes  an  area  which  is  referred  to 
the  Transition  life-zone,  but  which  has  a  ' '  Boreal  infusion. ' '  It  seems 
that  the  zones  of  life  found  on  the  upper  parts  of  southern  mountains 
show  less  affinity  to  particular  northern  transcontinental  life  belts 
than  they  do  to  an  alpine  or  Arctic  type  of  life  in  general. 

The  life-zones  of  the  various  parts  of  the  Sonoran  region  present 
still  greater  difficulties  in  homologizing.  In  the  first  place  it  may  be 
doubted  if  transcontinental  life-zones  really  show  in  the  best  manner 
the  similarities  and  differences  of  the  faunas  in  the  various  parts  of 
the  region.  Allen  (1892,  pp.  217-218)  has  demonstrated  that  the 
genera  and  subgenera  of  mammals  of  the  arid  division  of  the  Sonoran 
region  are  more  different  from  those  of  the  humid  division,  than  are 
those  of  a  northern  transcontinental  division  of  the  region  from  those 
of  a  southern  division.  It  seems,  then,  that  the  first  division  of  the 
Sonoran  region  should  be  into  eastern  and  western  sections. 

The  zones  of  the  Sonoran  region  found  in  the  various  parts  of  the 
western  United  States  are  very  difficult  to  correlate.  The  number  of 
zones  to  be  distinguished  is  variable  and  those  of  different  regions  do 
not  seem  to  be  exactly  homologous.  Following  Merriam  's  classification 
the  three  life-zones,  Transition,  Upper  Sonoran,  and  Lower  Sonoran 
have  usually  been  recognized.  However,  other  zones  are  sometimes 
apparent.  Grinnell  and  Swarth  (1913,  p.  217)  have  split  the  Transi- 
tion zone  in  the  San  Jacinto  area  of  southern  California  into  an  upper 
and  a  lower  division.  In  eastern  Washington  a  division  of  the  Transi- 
tion zone  has  also  been  made  and  these  divisions  are  as  distinct  as  are 
any  other  two  zones.  As  an  instance  of  the  difficulty  of  homologizing 
zones  in  different  regions,  we  may  mention  the  Columbia  Basin  prairie 


338  University  of  California  Publications  in  Zoology        [VOL.  16 

area.  This  area  seems  to  show  homologies  to  both  the  upper  Sonoran 
and  to  the  Transition  zones  as  found  in  other  parts  of  the  West. 

Certain  species  seem  to  have  a  different  "zonal"  position  in  dif- 
ferent regions.  Those  which  in  one  place  are  restricted  to  a  certain 
life-zone  range  elsewhere  into  areas  which  must  be  placed  in  other  life- 
zones.  Many  of  the  species  and  several  of  the  genera  given  by  Mer- 
riam  (1892  and  1898)  as  characteristic  of  the  various  life-zones  are 
now  known  to  range  beyond  the  limits  stated  by  him.  Grinnell  and 
Swarth  (1913,  p.  217)  mention  the  case  of  a  "Transition  infiltration 
into  a  prevailing  Upper  Sonoran  area"  in  the  San  Jacinto  region  of 
southern  California.  Cases  like  this  indicate  very  strongly  that  there 
is  often  a  lack  of  homology  between  the  zones  of  life  found  in  different 
regions. 

In  a  restricted  region  of  general  climatic  similarity  the  zones  of 
life  may  usually  be  easily  homologized.  In  California  Grinnell  (1902, 
p.  6)  has  recognized  several  zones  which  are  evidently  natural  divisions 
of  the  fauna,  and  each  of  which  is  seemingly  homologous  throughout 
its  extent  in  the  state.  However,  the  zones  of  life  found  in  different 
regions,  particularly  in  regions  under  different  climatic  conditions, 
show  much  less  similarity  and  in  many  cases  are  certainly  not  directly 
homologous. 

In  some  cases  the  life-zone  system  seems  to  be  largely  dependent 
upon  the  distribution  of  particular  associations  of  plants  and  animals. 
The  life-zones  are  based  on  temperature  differences,  yet  ' '  it  is  obvious 
that,  throughout  considerable  portions  of  the  continent,  the  details  of 
temperature  distribution  are  not  known  with  any  approach  to  pre- 
cision. Thus,  the  actual  criterion  which  the  field  zoologist  falls  back 
upon  in  any  given  case  is  the  character  of  the  fauna  and  flora  which 
he  finds  associated  together.  The  presence  of  certain  species  shows 
him  that  he  chances  to  be  in  this  or  that  'life-zone'  "  (Sumner,  1915, 
p.  67).  On  Alder  Creek  in  northern  Nevada,  Taylor  (1912,  p.  331) 
has  placed  the  vegetation  along  the  stream  in  the  Transition  zone, 
while  the  treeless  slopes  away  from  the  narrow  strip  of  vegetation  are 
placed  in  the  Upper  Austral  zone.  There  may  be  a  temperature  dif- 
ference between  the  strip  along  the  stream  and  the  immediately  ad- 
joining timberless  slopes  sufficiently  great  to  maintain  different  life- 
zones  in  the  two  places,  but  there  is  no  proof  that  such  is  the  case.  On 
the  contrary,  it  seems  that  the  differences  are  those  that  would  natur- 
ally be  produced  by  habitat  differences.  There  is  no  justification  for 
assuming  that  the  differences  in  this  and  many  other  similar  cases  are 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          339 

due  to  differences  in  temperature  until  the  effect  of  difference  in  habi- 
tat has  been  eliminated. 

We  believe  that  the  true  significance  of  the  facts  considered  in  the 
life-zone  method  of  studying  distribution  in  North  America  would  be 
better  presented  by  an  extension  of  the  zoogeographical  method  of 
Lydekker  and  Allen.  This  method  would  recognize  the  zones  of  life 
found  in  the  various  parts  of  the  continent.  It  would  recognize  the 
relation  of  the  zones  on  the  higher  parts  of  the  mountains  to_th_e  belts 
of  life  in  the  North  by  placing  these  higher  zones  in  subdivisions  of 
the  Holarctic  region.  Under  this  system  there  would  be  no  compulsion 
to  recognize  a  certain  number  of  life-zones  in  each  region,  but  the 
number  of  divisions  could  be  varied  to  fit  the  circumstances.  The 
effect  of  temperature  as  a  barrier  to  distribution  in  places  where  that 
is  important  would  be  shown,  and  the  effects  of  other  climatic  barriers 
could  also  be  emphasized. 


THE  ECOLOGICAL  METHOD 

The  ecological  method  of  studying  the  distribution  of  animals  and 
plants  brings  out  chiefly  the  relations  of  the  organisms  to  their  en- 
vironments. It  makes  as  simple  as  possible  the  comparison  of  environ- 
ments and  of  adaptational  structures  and  habits  in  different  species 
and  in  different  localities.  The  different  associations  and  formations 
of  any  region  can  be  compared  with  associations  and  formations  in  any 
other  part  of  the  world.  At  present  our  knowledge  of  associations  and 
formations  in  general  is  too  slight  to  point  out  the  significance  of  each 
ecological  division  in  southeastern  Washington,  but  we  feel  certain 
that  the  study  of  the  distribution  of  the  species  of  animals  in  relation 
to  the  distribution  of  different  kinds  of  environments  will  lead  to  re- 
sults of  the  highest  value. 

The  classification  of  the  habitats  of  a  region  and  the  placing  of  the 
species  in  associations,  which  to  some  extent  at  least  are  arbitrary 
divisions,  may  be  objected  to  on  the  ground  that  such  a  system  ap- 
parently indicates  a  discontinuity  in  nature  which  does  not  exist. 
However,  it  is  thought  that  the  use  of  terms  showing  the  relative 
abundance  of  each  species  in  the  different  habitats,  prevents  the  asso- 
ciations from  assuming  more  of  a  definite  character  than  they  actually 
possess. 

The  different  systems  used  for  describing  animal  distribution  are 
used  for  convenience  in  classifying  the  complexly  related  facts  in- 


340  University  of  California  Publications  in  Zoology        [VOL.  16 

volved.  It  seems  impossible  to  organize  all  the  facts  into  a  perfect 
system,  but  it  is  desirable  to  have  as  great  uniformity  as  possible  and 
not  unduly  to  increase  systems  or  complexity  of  nomenclature.  As 
the  facts  become  better  known  systems  will  have  to  be  changed  to  agree 
with  the  increased  knowledge.  At  present  it  seems  desirable  to  use 
two  systems,  both  starting  with  the  same  unit,  the  species.  By  the  first 
method,  zoogeography,  species  and  taxonomic  groups  are  considered 
in  relation  to  geographical  divisions.  The  second  method,  the  eco- 
logical method,  groups  species  according  to  similarity  of  adapt ational 
features  and  of  environmental  conditions. 

A  combination  of  these  two  methods  of  studying  distribution  should 
lead  to  excellent  results.  Usually  the  study  of  zoogeography  has  been 
carried  on  without  reference  to  the  particular  habitats  in  which  the 
organisms  live.  A  comparison  of  the  animals  in  similar  habitats  in 
different  faunal  areas  is  sure  to  bring  to  light  many  important  facts 
about  the  evolution  of  the  different  groups  and  of  topographical  and 
climatic  changes  in  general.  Further,  the  relation  between  an  organ- 
ism and  its  environment  cannot  be  fully  understood  without  reference 
to  the  mode  of  origin  both  of  the  organism  and  of  the  environment. 


SUMMARY 

In  southeastern  Washington  we  may  distinguish  three  faunal  areas, 
each  containing  a  number  of  distinct  habitats.  Each  habitat  is  occu- 
pied by  a  different  vertebrate  association. 

The  Columbia  Basin  sagebrush  fauna  belongs  to  the  Great  Basin 
division  of  the  Sonoran  region.  The  Blue  Mountain  fauna  belongs  to 
the  Canadian  subregion  of  Holarctica.  The  Columbia  Basin  prairie 
fauna  shows  affinities  to  the  life  both  of  the  Rocky  Mountains  and  of 
the  Great  Basin. 

An  Upper  Austral  life-zone,  a  timberless  and  a  timbered  division 
of  the  Transition  life-zone,  and  a  Boreal  life-zone  may  be  recognized 
in  the  region. 

Although  temperature  seems  to  be  the  climatic  barrier  which  is 
most  important  in  separating  the  faunas  of  the  zoogeographical 
regions  (Holarctic,  Sonoran,  and  Neotropical)  represented  in  America, 
yet  within  the  limits  of  the  Sonoran  region  it  has  not  been  proved  that 
temperature  is  as  important  a  barrier  to  distribution  as  are  the  factors 
connected  with  differences  in  rainfall  and  humidity. 


1916]        Dice:  Land  Vertebrates  of  Southeastern  Washington          341 

Zones  or  belts  of  life  may  be  recognized  in  many  regions.  How- 
ever, it  is  very  difficult  to  homologize  the  zones  of  life  which  occur  in 
widely  separated  parts  of  North  America.  The  division  of  the  conti- 
nent into  a  definite  number  of  transcontinental  life-zones  seems  to  be 
contrary  to  a  number  of  the  facts  of  distribution. 

The  ecological  method  of  studying  distribution  furnishes  valuable 
information  about  the  relation  between  organisms  and  their  environ- 
ments. The  use  of  this  method  in  conjunction  with  the  zoogeographi- 
cal  method  should  lead  to  results  of  great  value. 


LITERATURE  CITED 

ADAMS,  C.  C. 

1908.     The  ecological  succession  of  birds.    Auk,  25,  109-153. 
ALLEN,  J.  A. 

1892.     The  geographical   distribution   of   North  American   mammals.     Bull. 
Amer.  Mus.  Nat.  Hist.,  4,  199-244,  4  maps. 

BlGELOW,   F.   H. 

1908.     The  daily  normal  temperature  and  the  daily  normal  precipitation  of 
the  United  States.     U.  S.  Dept.  Agr.,  Weather  Bureau.  Bull.  E, 
186  pp. 
GADOW,  HANS. 

1913.  The  wanderings  of  animals  (University  Press,  Cambridge),  viii  -f-  150 

pp.,  17  maps. 
GATES,  FRANK  C. 

1911.     Summer  bird  life  in  the  vicinity  of  Havana,  Illinois,  in  its  relation  to 

the  prominent  plant  associations.    Wilson  Bull.,  23,  1-27. 
GRIN  NELL,  J. 

1902.     Check-list   of   California  birds.      Cooper  Ornith.   Club,   Pacific    Coast 
Avifauna,  3,  98  pp.,  2  col.  pis. 

1914.  An  account  of  the  mammals  and  birds  of  the  lower  Colorado  Valley 

with  especial  reference  to  the  distributional  problems  presented. 
Univ.  Calif.  Publ.  Zool.,  12,  51-294,  pis.  3-13,  9  figs. 
GRINNELL,  J.,  AND  SWARTH,  H.  S. 

1913.  An  account  of  the  birds  and  mammals  of  the  San  Jacinto  area  of 

southern    California,    with    remarks    upon    the    behavior    of    geo- 
graphic races  on  the  margins  of  their  habitats.     Univ.  Calif.  Publ. 
Zool.,  10,  197-406,  pis.  6-10,  3  figs. 
JACKSON,  HARTLEY  H.  T. 

1914.  The  land  vertebrates  of  Ridgway  Bog,  Wisconsin:   their  ecological 

succession  and  source  of  ingression.    Bull.  Wise.  Nat.  Hist.  Soc.,  12, 
4-54,  9  figs. 
KENNEDY,  C.  L. 

1914.     The  effects  of  irrigation  on  bird  life  in  the  Yakima  Valley,  Wash- 
ington.    Condor,  16,  250-255. 
LYDEKKER,  R. 

1896.     Geographical  history  of  mammals.      (University  Press,   Cambridge), 
xii  +  400  pp.,  82  figs.,  1  map. 


342  University  of  California  Publications  in  Zoology        [VOL.  16 

MERRIAM,  C.  HART. 

1890.  Eesults  of  a  biological  survey  of  the  San  Francisco  Mountain  region 

and  desert  of  the  Little  Colorado,  Arizona.  U.  S.  Dept.  AgrL,  Div. 
Orni.  and  Mamni.,  North  Amer.  Fauna,  3,  136  pp.,  14  pis.,  2  figs., 
5  maps. 

1891.  Eesults  of  a  biological  reconnoissance  of  southcentral  Idaho.     U.  S. 

Dept.  Agri.,  Div.  Orni.  and  Mamm.,  North  Amer.  Fauna,  5,  132 
pp.,  4  pis.,  4  figs. 

1892.  The  geographic   distribution  of  life  in  North  America  with  special 

reference  to  the  mammalia.    Proc.  Biol.  Soc.  Wash.,  7,  1-64,  map. 
1894.     Laws  of  temperature  control  of  the  geographic  distribution  of  ter- 

restial  animals  and  plants.     Natl.  Geog.  Mag.,  6,  229-238,  3  pis. 

(maps). 
1898.     Life  zones  and  crop  zones  in  the  United  States.     U.  S.  Dept.  Agri., 

Div.  Biol.  Surv.,  Bull.  No.  10,  79  pp.,  1  map. 
PIPER,  C.  V. 

1906.     Flora  of  the  State  of  Washington.    Smithsonian  Inst.,  Contrib.  U.  S. 

Natl.  Herb.,  11,  637  pp.,  22  pis.,  1  map. 

EUSSELL,  I.  C. 

1897.     A   reconnoissance   in   southeastern   Washington.     U.   S.   Geol.   Surv., 

Water  Supply  Paper  No.  4,  96  pp.,  7  pis.,  3  figs. 
SHELFORD,  V.  E. 

1911.  Physiological  animal  geography.     Jour.  Morph.,  22,  551-618,  19  figs. 

1913.  Animal    communities    in    temperate    America,    as    illustrated    in    the 

Chicago  region.  A  study  in  animal  ecology.  Bull.  Geog.  Soc.  Chi- 
cago, 5,  xiii  -f  362  pp.,  306  figs.,  2  maps. 

SUMNER,  F.  B 

1915.  Eeview  of  Grinnell  (1914):  An  account  of  the  mammals  and  birds  of 
the  lower  Colorado  Valley,  with  special  reference  to  the  distribu- 
tional problems  presented.  Science,  n.s.,  41,  65-69. 

TAYLOR,  W.  P. 

1912.  Field  notes  on  amphibians,  reptiles,  and  birds  of  northern  Humboldt 

County,  Nevada,  with  a  discussion  of  some  of  the  faunal  features 
of  the  region.    Univ.  Calif.  Publ.  Zool.,  7,  319-436,  pis.  7-12. 
WEAVER,  JOHN  E. 

1914.  Evaporation  and  plant   succession   in   southeastern   Washington   and 

adjacent  Idaho.    Plant  World,  17,  273-294,  10  figs. 


PLATE  24 

Fig.  1.  Packed  sand  situation  in  sagebrush  habitat,  three  miles  east  of 
Wallula,  June  13,  1914.  The  plants  are  common  sagebrush  (Artemisia  triden- 
tata)  and  rabbit  brush  (Chrysottiamnus  viscidiflorus  and  Chrysothamnus  nauseosus 
graveolens). 

Fig.  2.  Drifting  sand  situation  in  sagebrush  habitat,  three  miles  east  of 
Wallula,  June  13,  1914.  The  plants  are  common  sagebrush  and  rabbit  brush 
and  a  few  individuals  of  the  hop  sage  (  Grayia  spinosa).  In  the  distance  are 
the  bunchgrass-covered  hills  south  of  the  Walla  Walla  Kiver. 


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UNIV.   CALIF.    PUBL.   ZOOL.    VOL,    16 


[DICE]    PLATE  24 


PLATE  25 

Fig.  3.  Bunehgrass  habitat  in  the  prairie  area  one  mile  south  of  Lyon  's 
Ferry,  June  24,  1914.  The  principal  plant  is  the  wheat  bunchgrass  (Agropyron 
spicatum). 

Fig.  4.  The  Touchet  Valley,  two  miles  east  of  Prescott,  July  9,  1915.  The 
cottonwood-willow  habitat  is  shown  in  typical  development  along  the  river. 
The  conspicuous  trees  are  cotton  woods  (Populus  tricliocarpa} .  The  hills  and  the 
greater  part  of  the  valley  were  natively  covered  by  bunchgrass,  but  now  are 
nearly  entirely  in  cultivated  fields. 


[346] 


UNIV.   CALIF.    PUBL.  ZOOL.   VOL.    16 


[DICE]    PLATE  25 


Figure  3 


Figure  4 


PLATE  26 

Fig.  5.  The  eastern  side  of  the  canon  at  Hompeg  Falls,  Blue  Mountain  area. 
The  considerable  extent  of  the  rocky-slope  habitat  at  this  point  is  shown.  The 
trees  on  the  slopes  are  mostly  yellow  pines  (Pinus  ponderosa).  Toward  the 
upper  part  of  the  ridge  a  yellow-pine  habitat  occurs.  In  the  bottom  of  the 
canon  is  a  lowland-fir  habitat. 

Fig.  6.  Forests  in  the  Blue  Mountains  near  the  head  of  the  South  Fork  of 
the  Touchet  River,  Aug.  1,  1915.  The  forest  in  this  section  is  chiefly  made  up 
of  young  trees.  Douglas  spruce  (Pseudotsuga  taxi  folia)  and  western  larch 
(Larix  occidentalis)  are  the  dominant  species. 


[348] 


UNIV.   CALIF.    PUBL,  ZOOL.    VOL,   16 


[DICE]    PLATE  26 


f  * 


Figure  5 


Figure  6 


UNIVERSITY  OF  CALIFORNIA  PUBLICATIONS— (Continued) 

12.  Batrachoseps  major  and  Bufo  cognatus    calif ornicus,   New   Amphibia 

from  Southern  California,  by  Charles  Lewis  Camp.      Pp.  327-334. 
April,  1915 .10 

13.  Report  upon  Mammals  and  Birds  found  in  Portions  of  Trinity,  Siskiyou, 

and  Shasta  Counties,  California,  by  Louise  Kellogg.  Pp.  335-398, 
plates  15-18. 

14.  An  Analysis  of  the  Vertebrato  Fauna  of  the  Trinity  Region  of  Northern 

California,  by  Joseph  Grinnell.    Pp.  401-412. 

Nos.  13  and  14  in  one  cover.    January,  1916 75 

15.  The  Status  of  the  Beavers  of  Western  North  America,  with  a  Con- 

sideration of  the  Factors  in  their  Speciation,  by  Walter  P.  Taylor. 

Pp.  413-495,  22  text-figures.    March,  1916  85 

16.  Two  New  Aplodontias  from  Western  North  America,  by  Walter  P. 

Taylor.     Pp.  497-501.     May,  1916  .05 

Vol.  13.    1.  The  Schizopoda  of  the  San  Diego  Region,  by  Calvin  O.  Esterly.    Pp. 

1-20,  plates  1-2.    April,  1914 .15 

2.  A  Study  of  the  Occurrence  and  Manner  of  Distribution  of  the  Cteno- 
phora  of  the  San  Diego  Region,  by  Calvin  O.  Esterly.  Pp,  21-38. 
April,  1914  „ .15 

5.  A  New  Self -Regulating  Paraffin  Bath,  by  O.  W.  Woodworth.    Pp.  89- 

42,  2  text-figures.    April,  1914  .05 

4.  Diplodinium  ecaudatum,  with  an  Account  of  Its  Neuromotor  Apparatus, 
by  Robert  G.  Sharp.  Pp.  43-122,  plates  3-7,  4  text  figures.  May, 
1914 „_ 80 

6.  The  Vertical  Distribution  and  Movements  of  the  Schizopoda  of  the 

San  Diego  Region,  by  Calvin  O.  Esterly.    PD.  123-145.    May,  1914 .20 

6.  The  Anatomy  of  Heterodontus  francisci.     I.  The  Exoskeleton,  by  J. 

Frank  Daniel.  Pp.  147-166,  plates  8-9,  4  text  figures.  May  23, 
1914 _ „  .20 

7.  The  Movements  and  Reactions  of  the  Isolated  Melanophores  of  the 

Frog,  by  S.  J.  Holmes.    Pp.  167-174,  plate  10.    August,  1914 _      .10 

8.  Polychaetous  Annelids  of  the  Pacific  Coast  in  the  Collections  of  the 

Zoological  Museum  of  the  University  of  California,  by  Aaron  L. 
Treadwell.  Pp.  175-234,  plates  11-12. 

9.  New  Syllidae  from  San  Francisco  Bay    (collected  by  the  U.  S.  S. 

"Albatross"),  by  Aaron  L,  Treadwell.    Pp.  235-238,  7  text  figures. 
Nos.  8  and  9  in  one  cover.    October,  1914 65 

10.  Note  on  the  Medusan  Genus  Stomoloptius,  from  San  Diego,  by  Henry 

B.  Bigelow.     Pp.  239-241.     September,  1914  .05 

11.  A  Study  of  the  Structure  of  Feathers,  with  Reference  to  their  Taxo- 

nomic  Significance,  by  Asa  C.  Chandler.     Pp.  243-446,  plates  13-17, 

7  text-figures.    April,  1917 2.00 

12.  Anatomical  Adaptations  in  the  Thoracic  Limb  of  the  California  Pocket 

Gopher  and  Other  Rodents,  by  Charles  Daniel  Holliger.     Pp.  447- 

494,  plates  38-39,  20  text-figures.    March,  1916 45 

Vol.  14.  1.  A  Report  upon  the  Physical  Conditions  in  San  Francisco  Bay,  Based 
upon  the  Operations  of  the  United  States  Fisheries  Steamer  "Alba- 
tross" during  the  Years  1912  and  1913,  by  F.  B.  Sumner,  G.  D. 
Louderback,  W.  L.  Schmitt,  E.  C.  Johnston.  Pp.  1-198,  plates  1-13, 
20  text  figures.  July,  1914 2.25 

Vol.  15.  Introduction.  Dependence  of  Marine  Biology  upon  Hydrography  and 
Necessity  of  Quantitative  Biological  Research.  Pp.  i-xxiii.  June, 
1916 25 

1.  Hydrographic,  Plankton,  and  Dredging  Records  of  the  Scripps  Institu- 

tion for  Biological  Research  of  the  University  of  California,  1901  to 
1912,  compiled  and  arranged  under  the  supervision  of  W.  E.  Ritter 
by  Ellis  L.  Michael  and  George  F.  McEwen.  Pp.  1-206,  4  text  figures 
and  map.  July,  1915  , 2.25 

VoL  16.  1.  An  Outline  of  the  Morphology  and  Life  History  of  Crifhidia  lepto- 
coridis,  sp.  nov.,  by  Irene  McCulloch.  Pp.  1-22,  plates  1-4,  1  text 
figure.  September,  1915  „.  .25 

2.  On   Giardia  microti  sp.  nov.,   from  the  Meadow  Mouse,   by   Charles 

Atwood  Kofoid  and  Elizabeth  Bohn  Christiansen.  Pp.  23-29,  1  figure 
in  text. 

3.  On  Binary  and  Multiple  Fission  in  Giardia  muris  (Grassi),  by  Charles 

Atwood  Kofoid  and  Elizabeth  Bohn  Christiansen.  Pp.  30-54,  plates 
5-8,  1  figure  in  text. 

Nos.  2  and  3  in  one  cover.    November,  1915  ..  .30 


UNIVERSITY  OF  CALIFORNIA  PUBLICATIONS— (Continued) 

4.  The  Cultivation  of  Tissues  from  Amphibians,  by  John  C.  Johnson. 

Pp.  55-G2,  2  figures  in  text.    November,  1915  10 

5.  Notes  on  the  Tintinnoina.     1.  On  the  Probable  Origin  of  Dictyoeysta 

tiara  Haeckel.    2.  On  Petalotriclia  entzi  sp.  nov.,  by  Charles  Atwood 
Kof oid.    Pp.  63-69,  8  figures  in  text.    December,  1915 05 

6.  Binary  and  Multiple  Fission  in  Hexamitus,  by  Olive  Swezy.    Pp.  71-88, 

plates  9-11. 

7.  On  a  New  Trichomonad  Flagellate,  Tricliomitus  parvus,  from  the  Intes- 

tine of  Amphibians,  by  Olive  Swezy.    Pp.  89-94,  plate  12. 

Nos.  6  and  7  in  one  cover.    December,  1915 25 

8.  On  BlepJiarocorys  equi  sp.  nov.,  a  New  Ciliate  from  the  Caecum  of  the 

Horse,  by  Irwin  C.  Schumacher.    Pp.  95-106,  plate  13.     December, 

1915  _ 10 

9.  Three  New  Helices  from  California,  by  S.  Stillman  Berry.    Pp.  107- 

111.    January,  1916 05 

10.  On  Trypanosoma  triatomae,  a  New  Flagellate  from  a  Hemipteran  Bug 

from  the  Nests  of  the  Wood  Rat  Neotoma  fuscipes,  by  Charles  Atwood 
Kofoid  and  Irene  McCulloch.    Pp.  113-126,  plates  14-15.    February, 

1916 15 

11.  The  Genera  Monocercomonas  and  Polymastix,  by  Olive  Swezy.    Pp.  127- 

138,  plates  16-17.    February,  1916 10 

12.  Notes  on  the  Spiny  Lobster  (Panulirus  intermptus)  of  the  California 

Coast,  by  Bennet  M.  Allen.    Pp.  139-152,  2  figs,  in  text.    March,  1916      .15 

13.  Notes  on  the  Marine  Fishes  of  California,  by  Carl  L.  Hubbs.    Pp.  153- 

169,  plates  18-20.    March,  1916  15 

14.  The  Feeding  Habits  and  Food  of  Pelagic  Copepods  and  the  Question 

of  Nutrition  by  Organic  Substances  in  Solution  in  the  Water,  by 
Calvin  O.  Esterly.    Pp.  171-184,  2  figs,  in  text.    March,  1916 15 

15.  The  Kinetonucleus  of  Flagellates  and  the  Binuclear  Theory  of  Hart- 

mann,  by  Olive  Swezy.    Pp.  185-240,  58  figs,  in  text.    March,  1916 50 

16.  On  the  Life-History  of  a  Soil  Amoeba,  by  Charlie  Woodruff  Wilson. 

Pp.  241-292,  plates  18-23.     July,  1916  60 

17.  Distribution  of  the  Land  Vertebrates  of  Southeastern  Washington,  by 

Lee  Raymond  Dice.    Pp.  293-348,  plates  24-26.    June,  1916 60 


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